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Plant height affects directly the yield potential of wheat, and constitutes an important component of plant resistance to lodging. Although a large number of QTLs for plant height were detected, most of them have not been evaluated for their genetic effects yet. In the previous study, a major QTL Qph . nau-5B controlling wheat plant height was identified through association mapping in wheat variety collection. To evaluate the dwarfing effect of this QTL, three near-isogenic lines (NILs) with different alleles of Qph . nau-5B were developed using marker-assisted selection with Nanda 2419, Jichun 1016, and Zhengmai 9023 as donor and Zhongyou 9507 as receptor. The recipient genome compositions of these NILs were higher than 93%. Seven independent field trials were conducted and revealed that, compared with the recurrent parent, all NILs indicated a significant decrease in plant height (11.1 cm or 10.3% on average). Three alleles of Qph . nau-5B showed different degrees of dwarfing effects. The alleles coming from Jichun 1016 and Zhengmai 9023 displayed a similar effect on plant height (12.4 cm) in all environments, stronger than that of Nanda 2419 (8.6 cm). However, the relative dwarfism effects of different alleles were affected by different environments. Further analysis elucidated that this QTL had little detrimental influence on other agronomical traits such as spike number per plant, spike length and 1000-grain weight. These results suggested the breeding value of Qph . nau-5B that would be utilized for molecular design breeding of plant architecture in wheat.

KT1 was validated as a novel thickness QTL with major effects on wheat kernel dimensions and weight and fine mapped to a 0.04 cM interval near the chromosome-5A centromere. Kernel size, the principal grain weight determining factor of wheat and a target trait for both domestication and artificial breeding, is mainly defined by kernel length (KL), kernel width (KW) and kernel thickness (KT), of which KW and KT have been shown to be positively related to grain weight (GW). Qkt.nau-5A, a major QTL for KT, was validated using the QTL near-isogenic lines (NILs) in three genetic backgrounds. Genetic analysis using two F

Plant tissue brittleness is related to cellular structure and lodging. MED0031 is a mutant identified previously from ethyl methane sulfonate treatment of diploid wheat accession TA2726, showing brittleness in both stem and leaf. In microscopic and histological observations, the mutant was found to have less large vascular bundles per unit area, a thinner sclerenchyma cell wall, and a broader parenchyma, compared with the wild type. The mutated gene, TmBr1, was mapped to a 0.056 cM interval on chromosome 5Am. This gene was cloned using a MapRseq approach that searched the candidate gene through combination of the prior target gene mapping information with SNP calling and discovery of differentially expressed genes from RNA_seq data of the wild type and a BC3F2 bulk showing the mutant phenotype. TmBr1 encodes a COBL protein and a nonsense mutation within the region coding for the conserved COBRA domain caused premature translation termination. Introduction of TmBr1 to Arabidopsis AtCOBL4 mutant rescued the phenotype, demonstrating their functional conservation. Apart from the effect on cellulose content, the TmBr1 mutation might modulate synthesis of noncellulosic polysaccharide pectin as well. Application of the MapRseq approach to isolation of genes present in recombination cold spots and complicated genomes was discussed.

A major QTL QSpl.nau-7D, named HL2, was validated for its effects on head length and kernel number per spike using NIL, and mapped to a 0.2 cM interval using recombinants. Improvement in wheat inflorescence traits such as spike or head length and spikelet number provides an important avenue to increase grain yield potential. In a previous study, QSpl.nau-7D, the major QTL for head length on chromosome 7D, was identified in the recombinant inbred lines derived from Nanda2419 and Wangshuibai. To validate and precisely map this QTL, the Wangshuibai allele was transferred to elite cultivar Yangmai15 through marker-assisted selection. Compared with the recurrent parent, the resultant near-isogenic line (NIL) yielded not only 28% longer spikes on the average but also more spikelets and kernels per spike. Moreover, the NIL had a lower spikelet density and did not show significant kernel weight change. In the F2 population derived from the NIL, QSpl.nau-7D acted like a single semi-dominant gene controlling head length and was therefore designated as Head Length 2 (HL2). With this population, a high-density genetic map was constructed mainly using newly developed markers, and 100 homozygous recombinants including 17 genotypes were obtained. Field experiments showed that the recombinants carrying the 0.2-cM interval flanked by Xwgrb1588 and Xwgrb1902 from Wangshuibai produced longer spikes than those without this Wangshuibai allele. Comparative mapping of this interval revealed a conserved synteny among cereal grasses. HL2 is beneficial to wheat breeding for more kernels per spike at a lower spikelet density, which is a favored morphological trait for Fusarium head blight resistance.

Stably expressed type I and type II resistance QTL were identified using two Yangmai 158-derived RIL populations, and plant-height and flowering-time QTL intervals detected did not contribute to the FHB resistance variations. Yangmai 158 (Y158) is an elite wheat cultivar widely grown in China with stable Fusarium head blight (FHB) resistance. To enrich the genetic basis underlying FHB resistance, QTL mapping was conducted using two recombinant inbred line (RIL) populations derived from crosses of Y158 with susceptible lines Annong 8455 and Veery. Survey with makers linked to Fhb1, Fhb2, Fhb4 and Fhb5 in resistance cultivar Wangshuibai indicated that both Y158 and the susceptible lines do not contain these QTL. The RIL populations were surveyed with 65 PCR markers and 55 K chip, which generated 23,159 valid marker data, to produce genetic maps for whole genome scanning of quantitative trait loci (QTL). A total of six QTL, all with the Y158 alleles for better resistance and including one stably expressed QTL for type I resistance (Qfhi.nau-2D) and one stably expressed QTL for type II resistance (Qfhs.nau-2A), were identified. Moreover, taking advantage of the great genetic variations in plant height and flowering time, QTL conditioning these two traits were determined. Of six plant-height QTL and three flowering-time QTL intervals detected, none were associated with FHB resistance. The FHB resistance QTL in Y158 were shown to be useful alternatives in FHB resistance breeding programs. The SNP markers flanking Qfhs.nau-2A and Qfhi.nau-2D have been converted to breeder-friendly PCR-based markers to facilitate their applications.

Fine mapping of wheat powdery mildew-resistance gene Pm4e to a 0.19 cM interval with sequence-based markers provides the foundation for map-based cloning and marker-assisted selection with breeder-friendly markers. Powdery mildew caused by Blumeria graminis f. sp. tritici is a wheat foliar disease that poses a serious threat to global wheat production. Pm4 is a resistance gene locus that has played a key role in controlling this disease in wheat production and a few resistance alleles of this locus have been identified. We have previously mapped the Pm4e allele to a 6.7 cM interval on chromosome 2AL. In this study, Pm4e was delimited to a 0.19 cM interval flanked by Xwgrc763 and Xwgrc865, through employment of a larger segregating population, derived from the cross of resistant parent D29 with susceptible parent Yangmai 158 (Y158), and enrichment of the genetic interval with markers developed on Chinese Spring (C.S.) survey sequence. In this interval, Pm4e co-segregated with a few markers, some of which were either D29-dominant or Y158-dominant, implying great sequence variation in the interval between D29 and Y158. Most of these co-segregation markers could not differentiate the Pm4 alleles from each other. Survey of 55 wheat cultivars with four co-dominant markers showed that the Pm4e-co-segregating loci always co-exist. Annotation of the Pm4e interval-corresponding C.S. sequence revealed more than a dozen resistance gene analogs clustered in a 2.4 Mb region, although C.S. is susceptible to the Pm4e-avirulent isolate Bgt2. This study has established the foundation for map-based cloning of Pm4e. Moreover, some of the co-dominant markers developed in this study could help in marker-assisted transfer of Pm4e into elite cultivars.

Fusarium head blight (FHB), or scab, for its devastating nature to wheat production and food security, has stimulated worldwide attention. Multidisciplinary efforts have been made to fight against FHB for a long time, but the great progress has been achieved only in the genomics era of the past 20 years, particularly in the areas of resistance gene/QTL discovery, resistance mechanism elucidation and molecular breeding for better resistance. This review includes the following nine main sections, (1) FHB incidence, epidemic and impact, (2) causal Fusarium species, distribution and virulence, (3) types of host resistance to FHB, (4) germplasm exploitation for FHB resistance, (5) genetic control of FHB resistance, (6) fine mapping of Fhb1, Fhb2, Fhb4 and Fhb5, (7) cloning of Fhb1, (8) omics-based gene discovery and resistance mechanism study and (9) breeding for better FHB resistance. The advancements that have been made are outstanding and exciting; however, judged by the complicated nature of resistance to hemi-biotrophic pathogens like Fusarium species and lack of immune germplasm, it is still a long way to go to overcome FHB.

A major QTL QSpl.nau-7D, named HL2, was validated for its effects on head length and kernel number per spike using NIL, and mapped to a 0.2 cM interval using recombinants. Improvement in wheat inflorescence traits such as spike or head length and spikelet number provides an important avenue to increase grain yield potential. In a previous study, QSpl.nau-7D, the major QTL for head length on chromosome 7D, was identified in the recombinant inbred lines derived from Nanda2419 and Wangshuibai. To validate and precisely map this QTL, the Wangshuibai allele was transferred to elite cultivar Yangmai15 through marker-assisted selection. Compared with the recurrent parent, the resultant near-isogenic line (NIL) yielded not only 28% longer spikes on the average but also more spikelets and kernels per spike. Moreover, the NIL had a lower spikelet density and did not show significant kernel weight change. In the F

Unfortunately, the style of the units was incorrectly published (“cm” instead of “cM”) throughout the original article.

Head or ear blight, mainly caused by Fusarium species, can devastate almost all staple cereal crops (particularly wheat), resulting in great economic loss and imposing health threats on both human beings and livestock1-3. However, achievement in breeding for highly resistant cultivars is still not satisfactory. Here, we isolated the major-effect wheat quantitative trait locus, Qfhs.njau-3B, which confers head blight resistance, and showed that it is the same as the previously designated Fhb1. Fhb1 results from a rare deletion involving the 3' exon of the histidine-rich calcium-binding-protein gene on chromosome 3BS. Both wheat and Arabidopsis transformed with the Fhb1 sequence showed enhanced resistance to Fusarium graminearum spread. The translation products of this gene's homologs among plants are well conserved and might be essential for plant growth and development. Fhb1 could be useful not only for curbing Fusarium head blight in grain crops but also for improving other plants vulnerable to Fusarium species.

Wheat powdery mildew is a severe foliar disease and causes significant yield losses in epidemic years. Breeding and using resistant cultivars is the most widely employed strategy to curb this disease. To identify and transfer powdery mildew resistance genes in wild emmer wheat accession TA1410 into common wheat, a resistant F3 line derived from the cross of TA1410 × durum wheat line Zhongyin1320 was crossed with common wheat cultivar Yangmai158. The homozygous resistant BC5F2 lines derived from the backcross with Yangmai158 exhibited susceptibility at seedling stage and conferred increasing resistance when the plants were closer to heading stage. In two segregating BC5F3 families investigated at heading stage, the segregation of the resistance fit a 3:1 ratio, suggesting that a single dominant gene controls the resistance. This resistance gene, designated HSM1, was mapped to the 0.6-cM Xmag5825.1–Xgwm344 interval on chromosome 7AL and co-segregated with Xrga-C3 and Xrga-C6. A mapping position comparison with other powdery mildew resistance genes on this chromosome suggested that HSM1 belongs to the Pm1 resistance gene cluster. HSM1 is a useful candidate gene for resistance breeding, particularly in winter-wheat growing areas.

The spike characteristics length, spikelet density and fertile floret number are related yield components and are important in cereal improvement. QSpl.nau-2D is a major quantitative trait locus controlling spike length (SPL) detected in the recombinant inbred line population developed by crossing wheat (Triticum aestivum) cultivars Nanda2419 with Wangshuibai. In this study, to validate its genetic effect and determine its precise location, QSpl.nau-2D’s near-isogenic line (NIL) was developed using Mianyang99-323 as the recurrent parent through marker-assisted selection. Field trials showed that the NIL not only had significantly longer spikes on average than the recurrent parent but also had significantly higher grain weight, but did not differ in spikelet number and kernel number per spike. In the F2 population derived from a cross of the NIL with Mianyang99-323, QSpl.nau-2D functioned like a single gene and conditioned the SPL in a partially dominant manner, and was thus designated as HL1 (for head length). To precisely map HL1, 89 recombinants, consisting of 11 genotypes, were identified in the NIL-derived F2 population of 674 plants by using markers in the Xwmc25–Xgpw4080 interval. Phenotyping these lines showed that the introduction of a 0.9-cM interval flanked by Xcfd53 and DG371 in Nanda2419 resulted in longer spikes and a higher grain weight in the NIL. The availability of markers closely linked to HL1 could facilitate its use in breeding programs.

Understanding the genetics underlying yield formation of wheat is important for increasing wheat yield potential in breeding programs. Nanda2419 was a widely used cultivar for wheat production and breeding in China. In this study, we evaluated yield components and a few yield-related traits of a recombinant inbred line (RIL) population created by crossing Nanda2419 with the indigenous cultivar Wangshuibai in three to four trials at different geographical locations. Negative and positive correlations were found among some of these evaluated traits. Five traits had over 50 % trial-wide broad sense heritability. Using a framework marker map of the genome constructed with this population, quantitative trait loci (QTL) were identified for all traits, and epistatic loci were identified for seven of them. Our results confirmed some of the previously reported QTLs in wheat and identified several new ones, including QSn.nau-6D for effective tillers, QGn.nau-4B.2 for kernel number, QGw.nau-4D for kernel weight, QPh.nau-4B.2 and QPh.nau-4A for plant height, and QFlw.nau-5A.1 for flag leaf width. In the investigated population, Nanda2419 contributed all QTLs associated with higher kernel weight, higher leaf chlorophyll content, and a major QTL associated with wider flag leaf. Seven chromosome regions were related to more than one trait. Four QTL clusters contributed positively to breeding goal-based trait improvement through the Nanda2419 alleles and were detected in trials set in different ecological regions. The findings of this study are relevant to the molecular improvement of wheat yield and to the goal of screening cultivars for better breeding parents.

Flag leaf width (FLW) is directly related to photosynthetic capacity and yield potential in wheat. In a previous study, Qflw.nau-5A controlling FLW was detected on chromosome 5A in the interval possessing Fhb5 for type I Fusarium head blight (FHB) resistance using a recombinant inbred line population derived from Nanda2419 × Wangshuibai.Qflw.nau-5A near-isogenic line (NIL) with the background of Mianyang 99-323 and PH691 was developed and evaluated. FLW inheritance was investigated using two F2 populations developed from crossing the Qflw.nau-5A NILs with their recurrent parents. One hundred ten and 28 recombinants, which included 10 and 5 types of recombinants, were identified from 2816 F2 plants with Mianyang 99-323 background and 1277 F2 plants with PH691 background, respectively, and phenotyped in field trials for FLW and type I FHB resistance. Deletion bin mapping was applied to physically map Qflw.nau-5A.The introduction of Wangshuibai Qflw.nau-5A allele reduced the FLW up to 3 mm. In the F2 populations, Qflw.nau-5A was inherited like a semi-dominant gene, and was therefore designated as TaFLW1. The FLW of the recombinant lines displayed a distinct two-peak distribution. Recombinants with wider leaves commonly have Mianyang 99-323 or PH691 chromatin in the 0.2 cM Xwmc492-Xwmc752 interval that resided in the 5AL12-0.35-0.57 deletion bin, and recombinants with narrow leaves were Wangshuibai genotype in this interval. Phenotypic recombination between FLW and type I FHB resistance was identified, implying TaFLW1 was in close linkage with Fhb5. These results should aid wheat breeders to break the linkage drag through marker-assisted selection and assist in the map-based cloning of TaFLW1.

Seven kernel dimension QTLs were identified in wheat, and kernel thickness was found to be the most important dimension for grain weight improvement. Kernel morphology and weight of wheat (Triticum aestivum L.) affect both yield and quality; however, the genetic basis of these traits and their interactions has not been fully understood. In this study, to investigate the genetic factors affecting kernel morphology and the association of kernel morphology traits with kernel weight, kernel length (KL), width (KW) and thickness (KT) were evaluated, together with hundred-grain weight (HGW), in a recombinant inbred line population derived from Nanda2419 × Wangshuibai, with data from five trials (two different locations over 3 years). The results showed that HGW was more closely correlated with KT and KW than with KL. A whole genome scan revealed four QTLs for KL, one for KW and two for KT, distributed on five different chromosomes. Of them, QKl.nau-2D for KL, and QKt.nau-4B and QKt.nau-5A for KT were newly identified major QTLs for the respective traits, explaining up to 32.6 and 41.5% of the phenotypic variations, respectively. Increase of KW and KT and reduction of KL/KT and KW/KT ratios always resulted in significant higher grain weight. Lines combining the Nanda 2419 alleles of the 4B and 5A intervals had wider, thicker, rounder kernels and a 14% higher grain weight in the genotype-based analysis. A strong, negative linear relationship of the KW/KT ratio with grain weight was observed. It thus appears that kernel thickness is the most important kernel dimension factor in wheat improvement for higher yield. Mapping and marker identification of the kernel dimension-related QTLs definitely help realize the breeding goals.

Qfhi.nau-5A is a major quantitative trait locus (QTL) against Fusarium graminearum infection in the resistant wheat germplasm Wangshuibai. Genetic analysis using BC(3)F(2) and BC(4)F(2) populations, derived from selfing two near-isogenic lines (NIL) heterozygous at Qfhi.nau-5A that were developed, respectively, with Mianyang 99-323 and PH691 as the recurrent parent, showed that Qfhi.nau-5A inherited like a single dominant gene. This QTL was thus designated as Fhb5. To fine map it, these two backcross populations and a recombinant inbred line (RIL) population derived from Nanda2419 × Wangshuibai were screened for recombinants occurring between its two flanking markers Xbarc56 and Xbarc100. Nineteen NIL recombinants were identified from the two backcross populations and nine from the RIL population. In the RIL recombinant selection process, selection against Fhb4 present in the RIL population was incorporated. Genotyping these recombinant lines with ten markers mapping to the Xbarc56-Xbarc100 interval revealed four types of Mianyang 99-323-derived NIL recombinants, three types of PH691-derived NIL recombinants, and four types of RIL recombinants. In different field trials, the percentage of infected spikes of these lines displayed a distinct two-peak distribution. The more resistant class had over 55% less infection than the susceptible class. Common to these resistant genotypes, the 0.3-cM interval flanked by Xgwm304 and Xgwm415 or one of these two loci was derived from Wangshuibai, while none of the susceptible recombinants had Wangshuibai chromatin in this interval. This interval harboring Fhb5 was mapped to the pericentromeric C-5AS3-0.75 bin through deletion bin mapping. The precise localization of Fhb5 will facilitate its utilization in marker-assisted wheat breeding programs.

The characterization of yield trait mutants is important for understanding the regulation of grain yield formation in staple food crops. Meh0239 is a yield trait-related mutant identified from a mutant library of the common wheat cultivar Wangshuibai created by ethylmethyl sulfide (EMS) treatment of dry seeds. To shed some light on the nature of this mutation, it was investigated morphologically, physiologically, anatomically and genetically. The mutant plant showed obvious phenotypic differences in comparison with the wild type, starting at the seedling stage, including reduced plant height, wider and shorter leaves, shortened spikes, spikelets and grains and a more compact spikelet distribution. Also, seeds produced in the mutant germinated more slowly. Meh0239 contained a significantly higher level of abscisic acid (ABA) but lower levels of indole-3-acetic acid (IAA), methyl jasmonate (MeJA) and zeatin riboside (ZR) in flag leaves. Cells of all types in the leaf epidermis appeared shorter along the axial direction. The bulliform cells and long cells on the adaxial leaf surface were abnormal in shape. A genetic analysis using two F 2 segregating populations indicated that a single recessive mutation in wheat chromosome 7DS, about 3.1 cM distal from Xwmc506 , caused these variations. Because of the pleiotropic nature of this gene and its relation with yield trait formation, we named it Yt1 for yield trait related 1 .

Fusarium head blight or scab resistance in wheat is a complex quantitative trait affected greatly by environments. Therefore, the quantitative trait loci (QTL) for scab resistance found in mapping projects require validation to be effectively utilized in breeding programs. In this study, by employing both forward and background selections with the help of molecular markers, near-isogenic lines (NILs) for scab resistance QTLs Qfh.nau-2B, Qfhs.nau-3B, Qfhi.nau-4B and Qfhi.nau-5A, three of which originating in scab resistance germplasm Wangshuibai, were developed with the elite line Miangyang 99-323 as the recurrent parent. During the process of backcross, selection was based solely on marker genotypes of the target regions, and on recipient genome recovery rate in BC2F1 and BC3F1. All the identified BC3F1 plants with the target QTL regions have more than 94% recipient genome composition (RGC), and out of four to five of them a plant with over 97% RGC were obtained in each backcross combination. Compared with Mianyang 99-323, the Qfhs.nau-3B NIL showed much better resistance to disease spread within spikes, the Qfhi.nau-4B and Qfhi.nau-5A NILs showed much better resistance to initial infection, and the Qfh.nau-2B NIL showed improvement in both types of resistance. These results were consistent with findings in the previous QTL mapping studies. Morphologically and agronomically these NILs were similar to Mianyang 99-323 except that Qfhi.nau-4B NIL was taller and had a longer spike, and Qfhi.nau-5A NIL had narrower leaves. These results demonstrated the feasibility of marker-assisted utilization of scab resistance QTLs.

Bread wheat (Triticum aestivum L.) is a hexaploid species with a large and complex genome. A reference genetic marker map, namely the International Triticeae Mapping Initiative (ITMI) map, has been constructed with the recombinant inbred line population derived from a cross involving a synthetic line. But it is not sufficient for a full understanding of the wheat genome under artificial selection without comparing it with intervarietal maps. Using an intervarietal mapping population derived by crossing Nanda2419 and Wangshuibai, we constructed a high-density genetic map of wheat. The total map length was 4,223.1 cM, comprising 887 loci, 345 of which were detected by markers derived from expressed sequence tags (ESTs). Two-thirds of the high marker density blocks were present in interstitial and telomeric regions. The map covered, mostly with the EST-derived markers, approximately 158 cM of telomeric regions absent in the ITMI map. The regions of low marker density were largely conserved among cultivars and between homoeologous subgenomes. The loci showing skewed segregation displayed a clustered distribution along chromosomes and some of the segregation distortion regions (SDR) are conserved in different mapping populations. This map enriched with EST-derived markers is important for structure and function analysis of wheat genome as well as in wheat gene mapping, cloning, and breeding programs.

The QGw.nau - 2D, QGw.nau - 4B and QGw.nau - 5A intervals were investigated for their effects on weight, length, width, and thickness of kernels and their differential roles in determining kernel size and shape were demonstrated. Grain weight (GW) contributes greatly to wheat yield and is directly related to kernel size and shape. Although over 100 quantitative trait loci (QTLs) for GW have been reported in the literatures, few have been well characterized for their association with kernel traits. In this study, three GW QTLs identified in elite cultivar ‘Nanda2419’ (‘Mentana’), including QGw.nau-2D, QGw.nau-4B and QGw.nau-5A, were investigated through near isogenic line (NIL) development and evaluation. NILs for all three QTLs and one NIL with both QGw.nau-4B and QGw.nau-5A were developed with the help of marker-assisted selection after two to three generations of backcross using cultivar ‘Wangshuibai’ as the recurrent parent. One NIL with QGw.nau-4B in the background of cultivar ‘Wenmai6’ was also obtained. In four different field trials, these NILs consistently produced heavier kernels than the recurrent parents. QGw.nau-4B showed the largest effect on GW; its presence resulted in 0.4–0.5 g increase of hundred-grain weight, depending on genetic backgrounds. QGw.nau-4B and QGw.nau-5A functioned additively in conditioning GW. These three QTL intervals showed pleiotropic effects on, or close linkage with genes for, spike length, plant height and flag leaf width, respectively, and acted differentially in determining the kernel dimensions that are the major GW determinants. They all conditioned wider kernels with QGw.nau-5A displaying the largest effect. QGw.nau-4B and QGw.nau-5A also conditioned thicker kernels but had opposite effects on kernel length. This study demonstrated that marker-assisted selection is effective for GW improvement. The availability of GW NILs could facilitate cloning of GW genes and unraveling of kernel development mechanisms.

Powdery mildew, caused by Blumeria graminis f. sp. tritici (Bgt), is one of the most devastating foliar diseases of wheat and imposes a constant challenge on wheat breeders. Xiaohongpi, a Chinese landrace of wheat (Triticum aestivum L.), shows resistance to powdery mildew during the entire growth stage in the field and under controlled conditions. The F1 plants from cross of the powdery mildew susceptible cultivar Yangmai158 with Xiaohongpi were susceptible to isolate Bgt19, the locally most prevalent Bgt isolate. In the derived F2 population and F3 progenies, the resistance segregation deviated significantly from the one-gene Mendelian ratio. However, marker analysis indicated that only one recessive gene conferred the resistance, which co-segregated with Xsts-bcd1231 that showed co-segregation with Pm4a in different studies. Allelism test indicated that this recessive resistance gene, designated as pmX, is either allelic or tightly linked to Pm4a. The pmX gene was different from Pm4 alleles in resistance spectrum. Examination of the genotype frequencies at pmX and the linked marker loci in the F2 population showed that a genetic variation favoring the transmission of Xiaohongpi alleles could be the cause of deviated segregation. Mapping of the pmX-linked markers using Chinese Spring deletion lines indicated that it resides in the 0.85–1.00 bin of chromosome 2AL.

Powdery mildew, caused by Blumeria graminis f. sp. tritici, is one of the most serious wheat diseases. The rapid evolution of the pathogen's virulence, due to the heavy use of resistance genes, necessitates the expansion of resistance gene diversity. The common wheat line D57 is highly resistant to powdery mildew. A genetic analysis using an F(2) population derived from the cross of D57 with the susceptible cultivar Yangmai 158 and the derived F(2:3) lines indicated that D57 carries two dominant powdery mildew resistance genes. Based on mapping information of polymorphic markers identified by bulk segregant analysis, these two genes were assigned to chromosomes 5DS and 6DS. Using the F(2:3) lines that segregated in a single-gene mode, closely linked PCR-based markers were identified for both genes, and their chromosome assignments were confirmed through linkage mapping. The gene on chromosome 5DS was flanked by Xgwm205 and Xmag6176, with a genetic distance of 8.3 cM and 2.8 cM, respectively. This gene was 3.3 cM from a locus mapped by the STS marker MAG6137, converted from the RFLP marker BCD1871, which was 3.5 cM from Pm2. An evaluation with 15 pathogen isolates indicated that this gene and Pm2 were similar in their resistance spectra. The gene on chromosome 6DS was flanked by co-segregating Xcfd80 and Xmag6139 on one side and Xmag6140 on the other, with a genetic distance of 0.7 cM and 2.7 cM, respectively. This is the first powdery mildew resistance gene identified on chromosome 6DS, and plants that carried this gene were highly resistant to all of the 15 tested pathogen isolates. This gene was designated Pm45. The new resistance gene in D57 could easily be transferred to elite cultivars due to its common wheat origin and the availability of closely linked molecular markers.

Fusarium species cause serious diseases in cereal staple food crops such as wheat and maize. Currently, the mechanisms underlying resistance to Fusarium-caused diseases are still largely unknown. In the present study, we employed a combined proteomic and transcriptomic approach to investigate wheat genes responding to F. graminearum infection that causes Fusarium head blight (FHB). We found a total of 163 genes and 37 proteins that were induced by infection. These genes and proteins were associated with signaling pathways mediated by salicylic acid (SA), jasmonic acid (JA), ethylene (ET), calcium ions, phosphatidic acid (PA), as well as with reactive oxygen species (ROS) production and scavenging, antimicrobial compound synthesis, detoxification, and cell wall fortification. We compared the time-course expression profiles between FHB-resistant Wangshuibai plants and susceptible Meh0106 mutant plants of a selected set of genes that are critical to the plants' resistance and defense reactions. A biphasic phenomenon was observed during the first 24 h after inoculation (hai) in the resistant plants. The SA and Ca(2+) signaling pathways were activated within 6 hai followed by the JA mediated defense signaling activated around 12 hai. ET signaling was activated between these two phases. Genes for PA and ROS synthesis were induced during the SA and JA phases, respectively. The delayed activation of the SA defense pathway in the mutant was associated with its susceptibility. After F. graminearum infection, the endogenous contents of SA and JA in Wangshuibai and the mutant changed in a manner similar to the investigated genes corresponding to the individual pathways. A few genes for resistance-related cell modification and phytoalexin production were also identified. This study provided important clues for designing strategies to curb diseases caused by Fusarium.

Qfhi.nau-4B is a major quantitative trait locus (QTL) against Fusarium graminearum infection identified in the Fusarium head blight-resistant germplasm Wangshuibai. To fine map this QTL, a recombinant inbred line (RIL) population of 530 lines derived from Nanda2419 x Wangshuibai and the BC(3)F(2) population derived from the cross of a Qfhi.nau-4B near isogenic line (NIL) with susceptible cultivar Mianyang 99-323 as the recurrent parent were screened for recombinants occurred between microsatellite markers Xbarc20 and Xwmc349 that flank Qfhi.nau-4B. A total of 95 recombinants were obtained, including 45 RIL recombinants obtained through reverse-selection of Qfhi.nau-5A and 50 NIL recombinants from the BC(3)F(2) population. Genotyping these recombinant lines with 22 markers mapping to the Xbarc20 and Xwmc349 interval revealed fourteen genotypes of the RIL recombinants as well as of the NIL recombinants. Two-year field evaluation of their resistance to Fusarium infection showed that these lines could be clearly classified into two groups according to percentage of infected spikes. The more resistant class had over 60% less infection than the susceptible class and were common to have Wangshuibai chromatin in the 1.7-cM interval flanked by Xhbg226 and Xgwm149. None of the susceptible recombinants had this Wangshuibai chromatin. Qfhi.nau-4B was thus confined between Xhbg226 and Xgwm149 and named Fhb4. The interval harboring Fhb4 was mapped to 4BL5-0.86-1.00 bin using Chinese Spring deletion lines, a region with about 5.7 times higher recombination rate than the genome average. This study established the basis for map-based cloning of Fhb4.

Kernel number per spike is one of the most important yield components of wheat. To map QTLs related to kernel number including spike length (SPL), spikelet number per spike (SPN), fertile spikelet number (FSPN), sterile spikelet number (SSPN) and compactness, and to characterize the inheritance modes of the QTLs and two-locus interactions, 136 recombinant inbred lines (RILs) derived from 'Nanda2419' x 'Wangshuibai' and an immortalized F(2 )population (IF(2)) generated by randomly permutated intermating of these RILs were investigated. QTL mapping made use of the previously constructed over 3300 cM linkage map of the RIL population. Three, five, two, two and six chromosome regions were identified, respectively, for their association with SPL, SPN, FSPN, SSPN, and compactness in at least two of the three environments examined. All compactness QTLs but one shared the respective intervals of QSpn.nau-5A and the SPL QTLs. Xcfd46-Xwmc702 interval on chromosome 7D was related to all traits but SSPN and had consistently the largest effects. The fact that not all the compactness QTL intervals were related to both SPL and SPN indicates that compactness is regulated by different mechanisms. Interval coincidence between QTLs of SPL and SPN and between QTLs of FSPN and SSPN was minimal. For all the traits, favorable alleles exist in both parents. Inheritance modes from additiveness to overdominance of the QTLs were revealed and two-locus interactions were detected, implying that the traits studied are under complex genetic control. The results could contribute to wheat yield improvement and better use of Wangshuibai and Nanda2419 the two special germplasms in wheat breeding program.

Scab disease caused by Fusarium spp. has been a major concern for both wheat producers and consumers. Deployment of scab-resistant varieties is the major strategy to curb this disease. To identify the scab resistance genes in wheat cv. Wangshuibai, we produced a F(6:7) recombinant inbred line (RIL) population by crossing Wangshuibai with the scab-susceptible cultivar Nanda2419. The RILs were evaluated for scab resistance in the field by single floret inoculation in two replicates in 2002 and one replicate in 2003. The number of diseased spikelets (NDS) and the length of diseased rachides (LDR) were investigated to reflect the Type II resistance. Among 654 simple sequence repeat (SSR) markers surveyed, 326 were found to be polymorphic between the parents. A partial molecular map was constructed with these markers that covered over 2,210 cM of the wheat genome. Six chromosome regions showed association with both NDS and LDR in a one-way anova analysis, even though the variation explained by them varied between the two traits. Eight intervals were detected for their association with Type II resistance through interval mapping, five of which were not identified in single-point analysis. The quantitative trait loci (QTL) with large effects were the ones in the interval of Xgwm533-3-Xgwm533-1 on chromosome 3B and in the interval of Xwmc539-Xbarc024 on chromosome 6B, whose alleles favoring resistance originate from Wangshuibai. In addition, a QTL whose resistance allele originated from Nanda2419 was consistently detected in the interval of Xs1021m-Xgwm47-1 on chromosome 2B. These results suggest that Wangshuibai is the major source for Type II resistance in this population. The markers associated with these QTL would facilitate the use of scab-resistant genes of Wangshuibai in scab resistance breeding programs of wheat.