Recovery Plan for Wheat Blast Caused by Magnaporthe oryzae Pathotype Triticum

Wheat blast is an explosive new fungal disease of wheat caused by an Magnaporthe oryzae (synonym of Pyricularia oryzae) host-adapted subpopulation, the M. oryzae Triticum pathotype (MoT). MoT has been found in South America, South Asia, and Africa, but not in the United States. Wheat blast caused by the MoT fungus was first reported in Brazil in 1985 and subsequently spread to Bolivia, Paraguay, and Argentina in the 1990s and 2000s. The disease first appeared in Bangladesh in 2016 and in Zambia in 2017. The MoT fungus is seedborne, and the most likely route for movement across oceans was though grain trade. Wheat head (spike) blast is the predominant form of the disease in the field, although foliar and stem blast also occurs. The disease has proven hard to control when weather conditions are conducive, often resulting in devastating yield and quality losses. The only currently effective resistance, contained in the 2NvS translocation from the wild wheat relative Aegilops ventricosa, confers partial resistance that is variable depending on the genetic background of the specific wheat variety. Fungicides are not fully effective in controlling wheat head blast if warm, humid weather occurs during the heading stage. A major disease management strategy in areas where the disease occurs involves timing the wheat planting date so that heading does not coincide with warm rainy weather. A climate suitability model for the United States indicates that all of U.S. soft red winter wheat and about half of the hard red winter wheat are at risk.

Detection of breeding signatures in wheat using a linkage disequilibrium-corrected mapping approach

Marker assisted breeding, facilitated by reference genome assemblies, can help to produce cultivars adapted to changing environmental conditions. However, anomalous linkage disequilibrium (LD), where single markers show high LD with markers on other chromosomes but low LD with adjacent markers, is a serious impediment for genetic studies. We used a LD-correction approach to overcome these drawbacks, correcting the physical position of markers derived from 15 and 135 K arrays in a diversity panel of bread wheat representing 50 years of breeding history. We detected putative mismapping of 11.7% markers and improved the physical alignment of 5.4% markers. Population analysis indicated reduced genetic diversity over time as a result of breeding efforts. By analysis of outlier loci and allele frequency change over time we traced back the 2NS/2AS translocation of Aegilops ventricosa to one cultivar, “Cardos” (registered in 1998) which was the first among the panel to contain this translocation. A “selective sweep” for this important translocation region on chromosome 2AS was found, putatively linked to plant response to biotic stress factors. Our approach helps in overcoming the drawbacks of incorrectly anchored markers on the wheat reference assembly and facilitates detection of selective sweeps for important agronomic traits.

The Aegilops ventricosa 2NvS segment in bread wheat: cytology, genomics and breeding

Key message The first cytological characterization of the 2NvS segment in hexaploid wheat; complete de novo assembly and annotation of 2NvS segment; 2NvS frequency is increasing 2NvS and is associated with higher yield. Abstract The Aegilops ventricosa 2NvS translocation segment has been utilized in breeding disease-resistant wheat crops since the early 1990s. This segment is known to possess several important resistance genes against multiple wheat diseases including root knot nematode, stripe rust, leaf rust and stem rust. More recently, this segment has been associated with resistance to wheat blast, an emerging and devastating wheat disease in South America and Asia. To date, full characterization of the segment including its size, gene content and its association with grain yield is lacking. Here, we present a complete cytological and physical characterization of this agronomically important translocation in bread wheat. We de novo assembled the 2NvS segment in two wheat varieties, ‘Jagger’ and ‘CDC Stanley,’ and delineated the segment to be approximately 33 Mb. A total of 535 high-confidence genes were annotated within the 2NvS region, with > 10% belonging to the nucleotide-binding leucine-rich repeat (NLR) gene families. Identification of groups of NLR genes that are potentially N genome-specific and expressed in specific tissues can fast-track testing of candidate genes playing roles in various disease resistances. We also show the increasing frequency of 2NvS among spring and winter wheat breeding programs over two and a half decades, and the positive impact of 2NvS on wheat grain yield based on historical datasets. The significance of the 2NvS segment in wheat breeding due to resistance to multiple diseases and a positive impact on yield highlights the importance of understanding and characterizing the wheat pan-genome for better insights into molecular breeding for wheat improvement.

Genome-wide association mapping for wheat blast resistance in CIMMYT’s international screening nurseries evaluated in Bolivia and Bangladesh

Wheat blast caused by the fungus Magnaporthe oryzae pathotype Triticum (MoT) is an emerging threat to wheat production. To identify genomic regions associated with blast resistance against MoT isolates in Bolivia and Bangladesh, we performed a large genome-wide association mapping study using 8607 observations on 1106 lines from the International Maize and Wheat Improvement Centre’s International Bread Wheat Screening Nurseries (IBWSNs) and Semi-Arid Wheat Screening Nurseries (SAWSNs). We identified 36 significant markers on chromosomes 2AS, 3BL, 4AL and 7BL with consistent effects across panels or site-years, including 20 markers that were significant in all the 49 datasets and tagged the 2NS translocation from Aegilops ventricosa. The mean blast index of lines with and without the 2NS translocation was 2.7 ± 4.5 and 53.3 ± 15.9, respectively, that substantiates its strong effect on blast resistance. Furthermore, we fingerprinted a large panel of 4143 lines for the 2NS translocation that provided excellent insights into its frequency over years and indicated its presence in 94.1 and 93.7% of lines in the 2019 IBWSN and SAWSN, respectively. Overall, this study reinforces the effectiveness of the 2NS translocation for blast resistance and emphasizes the urgent need to identify novel non-2NS sources of blast resistance.

Resistance to Heterodera filipjevi and H. avenae in Winter Wheat is Conferred by Different QTL

The coexistence of cereal cyst nematode (CCN) species Heterodera avenae and H. filipjevi, often involving multiple pathotypes, is a limiting factor for wheat production in China. Some of the known genes for resistance to CCN are not effective against both nematode species, hence complicating breeding efforts to develop CCN-resistant wheat cultivars. Here, we demonstrate that the CCN resistance in wheat cultivar Madsen to both Heterodera spp. is controlled by different genetic loci, both of which originated from Aegilops ventricosa. A new quantitative trait locus (QTL), QCre-ma7D, was identified and localized in a 3.77-Mb genomic region on chromosome arm 7DL, which confers resistance to H. filipjevi. QCre-ma2A on chromosome arm 2AS corresponds to CCN resistance gene Cre5 and confers resistance to H. avenae. This QTL is a new locus on chromosome arm 7DL and is designated Cre9. Three Kompetitive allele-specific PCR markers (BS00150072, BS00021745, and BS00154302) were developed for molecular marker-assisted selection of Cre9 and locally adapted wheat lines with resistance to both nematode species were developed. QCre-ma2A on chromosome arm 2AS corresponds to CCN resistance gene Cre5 and confers resistance to H. avenae. The identification of different loci underlying resistance to H. filipjevi and H. avenae and the development of adapted resistant entries will facilitate breeding of wheat cultivars that are resistant to these devastating nematodes in China.

Novel Sources of Wheat Head Blast Resistance in Modern Breeding Lines and Wheat Wild Relatives

Wheat head blast (WHB), caused by the fungus Magnaporthe oryzae pathotype triticum, is a devastating disease affecting South America and South Asia. Despite 30 years of intensive effort, the 2NVS translocation from Aegilops ventricosa contains the only useful source of resistance to WHB effective against M. oryzae triticum isolates. The objective of this study was to identify non-2NVS sources of resistance to WHB among elite cultivars, breeding lines, landraces, and wild-relative accessions. Over 780 accessions were evaluated under field and greenhouse conditions in Bolivia, greenhouse conditions in Brazil, and at two biosafety level-3 laboratories in the United States. The M. oryzae triticum isolates B-71 (2012), 008 (2015), and 16MoT001 (2016) were used for controlled experiments, while isolate 008 was used for field experiments. Resistant and susceptible checks were included in all experiments. Under field conditions, susceptible spreaders were inoculated at the tillering stage to guarantee sufficient inoculum. Disease incidence and severity were evaluated as the average rating for each 1-m-row plot. Under controlled conditions, heads were inoculated after full emergence and individually rated for percentage of diseased spikelets. The diagnostic marker Ventriup-LN2 was used to test for the presence of the 2NVS translocation. Four non-2NVS spring wheat International Maize and Wheat Improvement Center breeding lines (CM22, CM49, CM52, and CM61) and four wheat wild-relatives (A. tauschii TA10142, TA1624, TA1667, and TA10140) were identified as resistant (<5% of severity) or moderately resistant (5 to <25% severity) to WHB. Experiments conducted at the seedling stage showed little correlation with disease severity at the head stage. M. oryzae triticum isolate 16MoT001 was significantly more aggressive against 2NVS-based varieties. The low frequency of WHB resistance and the increase in aggressiveness of newer M. oryzae triticum isolates highlight the threat that the disease poses to wheat production worldwide and the urgent need to identify and characterize new resistance genes that can be used in breeding for durably resistant varieties.

Yield reduction historically associated with the Aegilops ventricosa 7DV introgression is genetically and physically distinct from the eyespot resistance gene Pch1

Key message Yield penalty and increased grain protein content traits associated with Aegilops ventricosa 7D introgression have been mapped for the first time, and they are physically distinct from the eyespot resistance locus Pch1. Abstract Wheat wild relatives represent an important source of genetic variation, but introgression of agronomically relevant genes, such as for disease resistance, may lead to the simultaneous introduction of genetically linked deleterious traits. Pch1 is a dominant gene, conferring resistance to eyespot and was introgressed to wheat from Aegilops ventricosa as part of a large segment of the 7DV chromosome. This introgression has been associated with a significant yield reduction and a concomitant increase in grain protein content. In this study, we evaluated both traits and their relationship to the location of the Pch1 gene. We found that both QTLs were clearly distinct from the Pch1 gene, being located on a different linkage group to Pch1. In addition, we found that the QTL for increased grain protein content was strong and consistent across field trials, whereas the yield penalty QTL was unstable and environmentally dependent. The yield and grain protein content QTLs were genetically linked and located in the same linkage group. This finding is due in part to the small size of the population, and to the restricted recombination between wheat 7D and Ae. ventricosa 7Dv chromosomes. Although recombination in this interval is rare, it does occur. A recombinant line containing Pch1 and 7D_KASP6, the marker associated with increase in grain protein content, but not Xwmc221, the marker associated with the yield penalty effect, was identified.

How do eyespot resistance genes transferred into winter wheat breeding lines affect their yield?

Eyespot can reduce yields, even up to 50%. There are four genetically characterized resistances in wheat varieties, controlled by: (1) the Pch1 gene, transferred from Aegilops ventricosa; (2) the Pch2 gene, originating from wheat variety Capelle Desprez; (3) the Pch3 gene, originating from Dasypyrum villosum; and (4) the Q.Pch.jic-5A gene, a quantitative trait locus (QTL) located on chromosome 5A of Capelle Desprez. However, those loci have drawbacks, such as linkage of Pch1 with deleterious traits and limited effectiveness of Pch2 against the disease. Here we present an initial study which aims to characterize wheat pre-registration breeding lines carrying 12 eyespot resistance genes, consider their resistance expression in inoculation tests and the influence of resistance genotypes on the yield. We selected four groups of breeding lines, carrying: (1) the Pch1 gene alone: one line; (2) the Pch2 gene alone: four lines; (3) the Q.Pch.jic-5A gene alone: one line; and (4) Pch1 + Q.Pch.jic-5A: three lines. For the first time, the effect of the combination of Pch1 and Q.Pch.jic-5A genes was compared with resistance conferred by Pch1 or Q.Pch.jic-5A alone. We found significant differences between infection scores evaluated in resistant lines carrying Pch1 and Q.Pch.jic-5A alone, while no differences in terms of the level of resistance expression were detected between Pch1 alone and Pch1 + Q.Pch.jic-5A, and between wheat lines carrying Pch1 and Pch2 alone. Moreover, we demonstrated that the Pch1 gene, together with an Ae. ventricosa segment, caused statistically significant yield losses, both as a single eyespot resistance source or in a combination with Q.Pch.jic-5A. Yield scores showed that wheat lines with Q.Pch.jic-5A had the highest yields, similar to the yielding potential of Pch2-bearing lines and control varieties.