小麦遗传育种Wheat Genetics · Breeding

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Karyotype establishment and development of specific molecular markers of Aegilops geniculata Roth based on SLAF-seq

Yongfu Wang, Jianzhong Fan, Hong Zhang, Pingchuan Deng, Tingdong Li, Chunhuan Chen, Wanquan Ji, Yajuan Wang

2024, 23 (12): 3953-3965. DOI: 10.1016/j.jia.2023.09.014

Abstract （256） PDF in ScienceDirect

The constant evolution of pathogens poses a threat to wheat resistance against diseases, endangering food security. Developing resistant wheat varieties is the most practical approach for circumventing this problem. As a close relative of wheat, Aegilops geniculata, particularly accession SY159, has evolved numerous beneficial traits that could be applied to improve wheat. In this study, we established the karyotype of SY159 by fluorescence in situ hybridization (FISH) using the oligonucleotide probes Oligo-pTa535 and Oligo-pSc119.2 and a complete set of wheat–Ae. geniculata accession TA2899 addition lines as a reference. Using specific-locus amplified fragment sequencing (SLAF-seq) technology, 400 specific markers were established for detecting the SY159 chromosomes with efficiencies reaching 81.5%. The SY159-specific markers were used to classify the different homologous groups of SY159 against the wheat–Ae. geniculata addition lines. We used these specific markers on the 7M^g chromosome after classification, and successfully confirmed their suitability for studying the different chromosomes of SY159. This study provides a foundation for accelerating the application of SY159 in genetic breeding programs designed to improve wheat.

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Consensus linkage map construction and QTL mapping for eight yield-related traits in wheat using BAAFS 90K SNP array

Lihua Liu, Pingping Qu, Yue Zhou, Hongbo Li, Yangna Liu, Mingming Zhang, Liping Zhang, Changping Zhao, Shengquan Zhang, Binshuang Pang

2024, 23 (11): 3641-3656. DOI: 10.1016/j.jia.2023.07.028

Abstract （309） PDF in ScienceDirect

Identifying stable quantitative trait loci (QTLs) for yield-related traits across populations and environments is crucial for wheat breeding and genetic studies. Consensus maps also play important roles in wheat genetic and genomic research. In the present study, a wheat consensus map was constructed using a doubled haploid (DH) population derived from Jinghua 1×Xiaobaidongmai (JX), an F₂ population derived from L43×Shanxibaimai (LS) and the BAAFS Wheat 90K SNP array single nucleotide polymorphism (SNP) array. A total of 44,503 SNP markers were mapped on the constructed consensus map, and they covered 5,437.92 cM across 21 chromosomes. The consensus map showed high collinearity with the individual maps and the wheat reference genome IWGSC RefSeq v2.1. Phenotypic data on eight yield-related traits were collected in the JX population, as well as the F_2:3 and F_2:4 populations of LS, in six, two and two environments, respectively, and those data were used for QTL analysis. Inclusive composite interval mapping (ICIM) identified 32 environmentally stable QTLs for the eight yield-related traits. Among them, four QTLs (QPH.baafs-4B, QKNS.baafs-4B, QTGW.baafs-4B, and QSL.baafs-5A.3) were detected across mapping populations and environments, and nine stable QTLs (qKL.baafs-1D, QPH.baafs-2B, QKNS.baafs-3D, QSL.baafs-3D, QKW.baafs-4B, QPH.baafs-5D, QPH.baafs-6A.1, QSL.baafs-6A, and QSL.baafs-6D) are likely to be new. The physical region of 17.25–44.91 Mb on chromosome 4B was associated with six yield-related traits, so it is an important region for wheat yield. The physical region around the dwarfing gene Rht24 contained QTLs for kernel length (KL), kernel width (KW), spike length (SL), and thousand-grain weight (TGW), which are either from a pleiotropic effect of Rht24 or closely linked loci. For the stable QTLs, 254 promising candidate genes were identified. Among them, TraesCS5A03G1264300, TraesCS1B03G0624000 and TraesCS6A03G0697000 are particularly noteworthy since their homologous genes have similar functions for the corresponding traits. The constructed consensus map and the identified QTLs along with their candidate genes will facilitate the genetic dissection of wheat yield-related traits and accelerate the development of wheat cultivars with desirable plant morphology and high yield.

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Fine mapping and characterization of a major QTL for grain length, QGl.cau-2D.1, that has pleiotropic effects in synthetic allohexaploid wheat

Mingming Wang, Jia Geng, Zhe Zhang, Zihan Zhang, Lingfeng Miao, Tian Ma, Jiewen Xing, Baoyun Li, Qixin Sun, Yufeng Zhang, Zhongfu Ni

2024, 23 (9): 2911-2922. DOI: 10.1016/j.jia.2023.09.009

Abstract （151） PDF in ScienceDirect

Grain size is one of the determinants of grain yield, and identifying the genetic loci that control grain size will be helpful for increasing grain yield. In our previous study, a quantitative trait locus (QTL) for grain length (GL), QGl.cau-2D.1, was identified from an F2 population developed from the cross between the natural (TAA10) and synthetic (XX329) allohexaploid wheat. In the present study, we mainly fine mapped and validated its genetic effects. To this end, multiple near-isogenic lines (NILs) were obtained through marker-assisted selection with TAA10 as the recurrent parent. The secondary populations derived from 25 heterozygous recombinants were used for fine mapping of QGl.cau-2D.1, and the allele from XX329 significantly increased GL, thousand-grain weight (TGW), total spikelet number per spike (TSN) and spike compactness (SC). Using NILs for XX329 (2D+) and TAA10 (2D−), we determined the genetic and pleiotropic effects of QGl.cau-2D.1. The target sequences were aligned with the wheat reference genome RefSeq v2.1 and spanned an ~0.9 Mb genomic region. TraesCS2D03G0114900 (ortholog of Os03g0594700) was predicted as the candidate gene based on whole-genome re-sequencing and expression analyses. In summary, the map-based cloning of QGl.cau-2D.1 will be useful for improving grain weight with enhanced GL and TSN.

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Selection and application of four QTLs for grain protein content in modern wheat cultivars

Zihui Liu, Xiangjun Lai, Yijin Chen, Peng Zhao, Xiaoming Wang, Wanquan Ji, Shengbao Xu

2024, 23 (8): 2557-2570. DOI: 10.1016/j.jia.2023.09.006

Abstract （142） PDF in ScienceDirect

The grain protein content (GPC) is the key parameter for wheat grain nutritional quality. This study conducted a resampling GWAS analysis using 406 wheat accessions across eight environments, and identified four previously reported GPC QTLs. An analysis of 87 landraces and 259 modern cultivars revealed the loss of superior GPC haplotypes, especially in Chinese cultivars. These haplotypes were preferentially adopted in different agroecological zones and had broad effects on wheat yield and agronomic traits. Most GPC QTLs did not significantly reduce yield, suggesting that high GPC can be achieved without a yield penalty. The results of this study provide a reference for future GPC breeding in wheat using the four identified QTLs.

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Identification of P-type plasma membrane H⁺-ATPases in common wheat and characterization of TaHA7 associated with seed dormancy and germination

Bingli Jiang, Wei Gao, Yating Jiang, Shengnan Yan, Jiajia Cao, Litian Zhang, Yue Zhang, Jie Lu, Chuanxi Ma, Cheng Chang, Haiping Zhang

2024, 23 (7): 2164-2177. DOI: 10.1016/j.jia.2023.07.023

Abstract （170） PDF in ScienceDirect

The P-type plasma membrane (PM) H⁺-ATPases (HAs) are crucial for plant development, growth, and defense. The HAs have been thoroughly characterized in many different plants. However, despite their importance, the functions of HAs in germination and seed dormancy (SD) have not been validated in wheat. Here, we identified 28 TaHA genes (TaHA1-28) in common wheat, which were divided into five subfamilies. An examination of gene expression in strong- and weak-SD wheat varieties led to the discovery of six candidate genes (TaHA7/-12/-14/-16/-18/-20). Based on a single nucleotide polymorphism (SNP) mutation (C/T) in the TaHA7 coding region, a CAPS marker (HA7) was developed and validated in 168 wheat varieties and 171 Chinese mini-core collections that exhibit diverse germination and SD phenotypes. We further verified the roles of the two allelic variations of TaHA7 in germination and SD using wheat mutants mutagenized with ethyl methane sulphonate (EMS) in ‘Jimai 22’ and ‘Jing 411’ backgrounds, and in transgenic Arabidopsis lines. TaHA7 appears to regulate germination and SD by mediating gibberellic acid (GA) and abscisic acid (ABA) signaling, metabolism, and biosynthesis. The results presented here will enable future research regarding the TaHAs in wheat.

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Establishment of a transformation system in close relatives of wheat under the assistance of TaWOX5

Yanan Chang, Junxian Liu, Chang Liu, Huiyun Liu, Huali Tang, Yuliang Qiu, Zhishan Lin, Ke Wang, Yueming Yan, Xingguo Ye

2024, 23 (6): 1839-1849. DOI: 10.1016/j.jia.2023.06.021

Abstract （263） PDF in ScienceDirect

Species closely related to wheat are important genetic resources for agricultural production, functional genomics studies and wheat improvement. In this study, a wheat gene related to regeneration, TaWOX5, was applied to establish the Agrobacterium-mediated transformation systems of Triticum monococcum, hexaploid triticale, and rye (Secale cereale L.) using their immature embryos. Transgenic plants were efficiently generated. During the transformation process, the Agrobacterium infection efficiency was assessed by histochemical staining for β-glucuronidase (GUS). Finally, the transgenic nature of regenerated plants was verified by polymerase chain reaction (PCR)-based genotyping for the presence of the GUS and bialaphos resistance (bar) genes, histochemical staining for GUS protein, and the QuickStix strip assay for bar protein. The transformation efficiency of T. monococcum genotype PI428182 was 94.4%; the efficiencies of four hexaploid triticale genotypes Lin456, ZS3297, ZS1257, and ZS3224 were 52.1, 41.2, 19.4, and 16.0%, respectively; and the transformation efficiency of rye cultivar Lanzhou Heimai was 7.8%. Fluorescence in situ hybridization (FISH) and genomic in situ hybridization (GISH) analyses indicated that the GUS transgenes were integrated into the distal or near centromere (proximal) regions of the chromosomes in transgenic T. monococcum and hexaploid triticale plants. In the transgenic hexaploid triticale plants, the foreign DNA fragment was randomly integrated into the AABB and RR genomes. Furthermore, the transgene was almost stably inherited in the next generation by Mendel’s law. The findings in this study will promote the genetic improvement of the three plant species for grain or forage production and the improvement of cereal species including wheat for functional genomics studies.

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Genome-wide association study of grain micronutrient concentrations in bread wheat

Yongchao Hao, Fanmei Kong, Lili Wang, Yu Zhao, Mengyao Li, Naixiu Che, Shuang Li, Min Wang, Ming Hao, Xiaocun Zhang, Yan Zhao

2024, 23 (5): 1468-1480. DOI: 10.1016/j.jia.2023.06.030

Abstract （251） PDF in ScienceDirect

Bread wheat (Triticum aestivum) is a staple food crop worldwide. The genetic dissection of important nutrient traits is essential for the biofortification of wheat to meet the nutritional needs of the world’s growing population. Here, 45,298 single-nucleotide polymorphisms (SNPs) from 55K chip arrays were used to genotype a panel of 768 wheat cultivars, and a total of 154 quantitative trait loci (QTLs) were detected for eight traits under three environments by genome-wide association study (GWAS). Three QTLs (qMn-3B.1, qFe-3B.4, and qSe-3B.1/qFe-3B.6) detected repeatedly under different environments or traits were subjected to subsequent analyses based on linkage disequilibrium decay and the P-values of significant SNPs. Significant SNPs in the three QTL regions formed six haplotypes for qMn-3B.1, three haplotypes for qFe-3B.4, and three haplotypes for qSe-3B.1/qFe-3B.6. Phenotypic analysis revealed significant differences among haplotypes. These results indicated that the concentrations of several nutrient elements have been modified during the domestication of landraces to modern wheat. Based on the QTL regions, we identified 15 high-confidence genes, eight of which were stably expressed in different tissues and/or developmental stages. TraesCS3B02G046100 in qMn-3B.1 and TraesCS3B02G199500 in qSe-3B.1/qFe-3B.6 were both inferred to interact with metal ions according to the Gene Ontology (GO) analysis. TraesCS3B02G199000, which belongs to qSe-3B.1/qFe-3B.6, was determined to be a member of the WRKY gene family. Overall, this study provides several reliable QTLs that may significantly affect the concentrations of nutrient elements in wheat grain, and this information will facilitate the breeding of wheat cultivars with improved grain properties.

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Transcriptomic and metabolomic analysis provides insights into lignin biosynthesis and accumulation and differences in lodging resistance in hybrid wheat

YANG Wei-bing, ZHANG Sheng-quan, HOU Qi-ling, GAO Jian-gang, WANG Han-Xia, CHEN Xian-Chao, LIAO Xiang-zheng, ZHANG Feng-ting, ZHAO Chang-ping, QIN Zhi-lie

2024, 23 (4): 1105-1117. DOI: 10.1016/j.jia.2023.06.027

Abstract （230） PDF in ScienceDirect

The use of hybrid wheat is one way to improve the yield in the future. However, greater plant heights increase lodging risk to some extent. In this study, two hybrid combinations with differences in lodging resistance were used to analyze the stem-related traits during the filling stage, and to investigate the mechanism of the difference in lodging resistance by analyzing lignin synthesis of the basal second internode (BSI). The stem-related traits such as the breaking strength, stem pole substantial degree (SPSD), and rind penetration strength (RPS), as well as the lignin content of the lodging-resistant combination (LRC), were significantly higher than those of the lodging-sensitive combination (LSC). The phenylpropanoid biosynthesis pathway was significantly and simultaneously enriched according to the transcriptomics and metabolomics analysis at the later filling stage. A total of 35 critical regulatory genes involved in the phenylpropanoid pathway were identified. Moreover, 42% of the identified genes were significantly and differentially expressed at the later grain-filling stage between the two combinations, among which more than 80% were strongly up-regulated at that stage in the LRC compared with LSC. On the contrary, the LRC displayed lower contents of lignin intermediate metabolites than the LSC. These results suggested that the key to the lodging resistance formation of LRC is largely the higher lignin synthesis at the later grain-filling stage. Finally, breeding strategies for synergistically improving plant height and lodging resistance of hybrid wheat were put forward by comparing the LRC with the conventional wheat applied in large areas.

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Creating large EMS populations for functional genomics and breeding in wheat

Wenqiang Wang, Xizhen Guan, Yong Gan, Guojun Liu, Chunhao Zou, Weikang Wang, Jifa Zhang, Huifei Zhang, Qunqun Hao, Fei Ni, Jiajie Wu, Lynn Epstein, Daolin Fu

2024, 23 (2): 484-493. DOI: 10.1016/j.jia.2023.05.039

Abstract （592） PDF in ScienceDirect

Wheat germplasm is a fundamental resource for basic research, applied studies, and wheat breeding, which can be enriched normally by several paths, such as collecting natural lines, accumulating breeding lines, and introducing mutagenesis materials. Ethyl methane sulfonate (EMS) is an alkylating agent that can effectively introduce genetic variations in a wide variety of plant species. In this study, we created a million-scale EMS population (MEP) that started with the Chinese wheat cultivars ‘Luyan 128’, ‘Jimai 38’, ‘Jimai 44’, and ‘Shannong 30’. In the M₁ generation, the MEP had numerous phenotypical variations, such as >3,000 chlorophyll-deficient mutants, 2,519 compact spikes, and 1,692 male sterile spikes. There were also rare mutations, including 30 independent tillers each with double heads. Some M₁ variations of chlorophyll-deficiency and compact spikes were inheritable, appearing in the M₂ or M₃ generations. To advance the entire MEP to higher generations, we adopted a single-seed descendent (SSD) approach. All other seed composites of M₂ were used to screen other agronomically important traits, such as the tolerance to herbicide quizalofop-P-methyl. The MEP is available for collaborative projects, and provides a valuable toolbox for wheat genetics and breeding for sustainable agriculture.

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Assessment of molecular markers and marker-assisted selection for drought tolerance in barley (Hordeum vulgare L.)

Akmaral Baidyussen, Gulmira Khassanova, Maral Utebayev, Satyvaldy Jatayev, Rystay Kushanova, Sholpan Khalbayeva, Aigul Amangeldiyeva, Raushan Yerzhebayeva, Kulpash Bulatova, Carly Schramm, Peter Anderson, Colin L. D. Jenkins, Kathleen L. Soole, Yuri Shavrukov

2024, 23 (1): 20-38. DOI: 10.1016/j.jia.2023.06.012

Abstract （152） PDF in ScienceDirect

This review updates the present status of the field of molecular markers and marker-assisted selection (MAS), using the example of drought tolerance in barley. The accuracy of selected quantitative trait loci (QTLs), candidate genes and suggested markers was assessed in the barley genome cv. Morex. Six common strategies are described for molecular marker development, candidate gene identification and verification, and their possible applications in MAS to improve the grain yield and yield components in barley under drought stress. These strategies are based on the following five principles: (1) Molecular markers are designated as genomic ‘tags’, and their ‘prediction’ is strongly dependent on their distance from a candidate gene on genetic or physical maps; (2) plants react differently under favourable and stressful conditions or depending on their stage of development; (3) each candidate gene must be verified by confirming its expression in the relevant conditions, e.g., drought; (4) the molecular marker identified must be validated for MAS for tolerance to drought stress and improved grain yield; and (5) the small number of molecular markers realized for MAS in breeding, from among the many studies targeting candidate genes, can be explained by the complex nature of drought stress, and multiple stress-responsive genes in each barley genotype that are expressed differentially depending on many other factors.

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Genome-wide association and linkage mapping strategies reveal the genetic loci and candidate genes of important agronomic traits in Sichuan wheat

ZHANG Zhi-peng, LI Zhen, HE Fang, LÜ Ji-juan, XIE Bin, YI Xiao-yu, LI Jia-min, LI Jing, SONG Jing-han, PU Zhi-en, MA Jian, PENG Yuan-ying, CHEN Guo-yue, WEI Yu-ming, ZHENG You-liang, LI Wei

2023, 22 (11): 3380-3393. DOI: 10.1016/j.jia.2023.02.030

Abstract （238） PDF in ScienceDirect

Increasing wheat yield is a long-term goal for wheat breeders around the world. Exploiting elite genetic resources and dissecting the genetic basis of important agronomic traits in wheat are the necessary methods for high-yield wheat breeding. This study evaluated nine crucial agronomic traits found in a natural population of 156 wheat varieties and 77 landraces from Sichuan, China in seven environments over two years. The results of this investigation of agronomic traits showed that the landraces had more tillers and higher kernel numbers per spike (KNS), while the breeding varieties had higher thousand-kernel weight (TKW) and kernel weight per spike (KWS). The generalized heritability (H²) values of the nine agronomic traits varied from 0.74 to 0.95. Structure analysis suggested that the natural population could be divided into three groups using 43 198 single nucleotide polymorphism (SNP) markers from the wheat 55K SNP chip. A total of 67 quantitative trait loci (QTLs) were identified by the genome-wide association study (GWAS) analysis based on the Q+K method of a mixed linear model. Three important QTLs were analyzed in this study. Four haplotypes of QFTN.sicau-7BL.1 for fertile tillers number (FTN), three haplotypes of QKNS.sicau-1AL.2 for KNS, and four haplotypes of QTKW.sicau-3BS.1 for TKW were detected. FTN-Hap2, KNS-Hap1, and TKW-Hap2 were excellent haplotypes for each QTL based on the yield performance of 42 varieties in regional trials from 2002 to 2013. The varieties with all three haplotypes showed the highest yield compared to those with either two haplotypes or one haplotype. In addition, the KASP-AX-108866053 marker of QTL QKNS.sicau-1AL.2 was successfully distinguished between three haplotypes (or alleles) in 63 varieties based on the number of kernels per spike in regional trials between 2018 and 2021. These genetic loci and reliable makers can be applied in marker-assisted selection or map-based gene cloning for the genetic improvement of wheat yield.

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Rapid identification of Psathyrostachys huashanica Keng chromosomes in wheat background based on ND-FISH and SNP array methods

LI Jia-chuang, LI Jiao-jiao, ZHAO Li, ZHAO Ji-xin, WU Jun, CHEN Xin-hong, ZHANG Li-yu, DONG Pu-hui, WANG Li-ming, ZHAO De-hui, WANG Chun-ping, PANG Yu-hui

2023, 22 (10): 2934-2948. DOI: 10.1016/j.jia.2023.02.001

Abstract （242） PDF in ScienceDirect

Psathyrostachys huashanica Keng (2n=2x=14, NsNs) is regarded as a valuable wild relative species for common wheat cultivar improvement because of its abundant beneficial agronomic traits. However, although the development of many wheat–P. huashanica-derived lines provides a germplasm base for the transfer of excellent traits, the lag in the identification of P. huashanica chromosomes in the wheat background has limited the study of these lines. In this study, three novel nondenaturing fluorescence in situ hybridization (ND-FISH)-positive oligo probes were developed. Among them, HS-TZ3 and HS-TZ4 could specifically hybridize with P. huashanica chromosomes, mainly in the telomere area, and HS-CHTZ5 could hybridize with the chromosomal centromere area. We sequentially constructed a P. huashanica FISH karyotype and idiogram that helped identify the homologous groups of introduced P. huashanica chromosomes. In detail, 1Ns and 2Ns had opposite signals on the short and long arms, 3Ns, 4Ns, and 7Ns had superposed two-color signals, 5Ns and 6Ns had fluorescent signals only on their short arms, and 7Ns had signals on the intercalary of the long arm. In addition, we evaluated different ways to identify alien introgression lines by using low-density single nucleotide polymorphism (SNP) arrays and recommended the SNP homozygosity rate in each chromosome as a statistical pattern. The 15K SNP array is widely applicable for addition, substitution, and translocation lines, and the 40K SNP array is the most accurate for recognizing transposed intervals between wheat and alien chromosomes. Our research provided convenient methods to distinguish the homologous group of P. huashanica chromosomes in a common wheat background based on ND-FISH and SNP arrays, which is of great significance for efficiently identifying wheat–P. huashanica-derived lines and the further application of Ns chromosomes

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SNP-based identification of QTLs for thousand-grain weight and related traits in wheat 8762/Keyi 5214 DH lines

HUANG Feng, LI Xuan-shuang, DU Xiao-yu, LI Shun-cheng, LI Nan-nan, LÜ Yong-jun, ZOU Shao-kui, ZHANG Qian, WANG Li-na, NI Zhong-fu, HAN Yu-lin, XING Jie-wen

2023, 22 (10): 2949-2960. DOI: 10.1016/j.jia.2023.03.004

Abstract （312） PDF in ScienceDirect

As important yield-related traits, thousand-grain weight (TGW), grain number per spike (GNS) and grain weight per spike (GWS) are crucial components of wheat production. To dissect their underlying genetic basis, a double haploid (DH) population comprised of 198 lines derived from 8762/Keyi 5214 was constructed. We then used genechip to genotype the DH population and integrated the yield-related traits TGW, GNS and GWS for QTL mapping. Finally, we obtained a total of 18 942 polymorphic SNP markers and identified 41 crucial QTLs for these traits. Three stable QTLs for TGW were identified on chromosomes 2D (QTgw-2D.3 and QTgw-2D.4) and 6A (QTgw-6A.1), with additive alleles all from the parent 8762, explaining 4.81–18.67% of the phenotypic variations. Five stable QTLs for GNS on chromosomes 3D, 5B, 5D and 6A were identified. QGns-5D.1 was from parent 8762, while the other four QTLs were from parent Keyi 5214, explaining 5.89–7.08% of the GNS phenotypic variations. In addition, a stable GWS genetic locus QGws-4A.3 was detected from the parent 8762, which explained 6.08–6.14% of the phenotypic variations. To utilize the identified QTLs, we developed STARP markers for four important QTLs, Tgw2D.3-2, Tgw2D.4-1, Tgw6A.1 and Gns3D.1. Our results provide important basic resources and references for the identification and cloning of genes related to TGW, GNS and GWS in wheat.

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Dissecting the key genomic regions underlying high yield potential in common wheat variety ‘Kenong 9204’

ZHAO Chun-hua, ZHANG Na, FAN Xiao-li, JI Jun, SHI Xiao-li, CUI Fa, LING Hong-qing, LI Jun-ming

2023, 22 (9): 2603-2616. DOI: 10.1016/j.jia.2023.02.013

Abstract （330） PDF in ScienceDirect

The foundation parents play key roles in the genetic improvement of both yield potential and end-use quality in wheat. Characterizing the genetic basis that underlies certain beneficial traits in the foundation parents will provide theoretical reference for molecular breeding by a design approach. ‘Kenong 9204’ (KN9204) is a candidate foundation parent characterized by ideotype, high yield potential, and particularly high nitrogen fertilizer utilization. To better understand the genetic basis of its high yield potential, high throughput whole-genome re-sequencing (10×) was performed on KN9204, its parental lines and its derivatives. A high-resolution genetic composition map of KN9204 was constructed, which showed the parental origin of the favorable genomic segments based on the identification of excellent yield-related quantitative trait loci (QTL) from a bi-parental mapping population. Xiaoyan 693 (XY693), a wheat–Thinopyrum ponticum partial amphidiploid, contributed a great deal to the high yield potential of KN9204, and three major stable QTLs from XY693 were fine mapped. The transmissibility of key genomic segments from KN9204 to its derivatives were delineated, indicating that haplotype blocks containing beneficial gene combinations were conserved along with directional selection by breeders. Evidence for selection sweeps in the breeding programs was identified. This study provides a theoretical reference for the breeding of high-yield wheat varieties by a molecular design approach.

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Dissecting the genetic basis of grain color and pre-harvest sprouting resistance in common wheat by association analysis

YAN Sheng-nan, YU Zhao-yu, GAO Wei, WANG Xu-yang, CAO Jia-jia, LU Jie, MA Chuan-xi, CHANG Cheng, ZHANG Hai-ping

2023, 22 (9): 2617-2631. DOI: 10.1016/j.jia.2023.04.017

Abstract （262） PDF in ScienceDirect

Pre-harvest sprouting (PHS) adversely affects wheat quality and yield, and grain color (GC) is associated with PHS resistance. However, the genetic relationship between GC and PHS resistance remains unclear. In this study, 168 wheat varieties (lines) with significant differences in GC and PHS resistance were genotyped using an Illumina 90K iSelect SNP array. Genome-wide association study (GWAS) based on a mixed linear model showed that 67 marker-trait associations (MTAs) assigned to 29 loci, including 17 potentially novel loci, were significantly associated with GC, which explained 1.1–17.0% of the phenotypic variation. In addition, 100 MTAs belonging to 54 loci, including 31 novel loci, were significantly associated with PHS resistance, which accounted for 1.1–14.7% of the phenotypic variation. Subsequently, two cleaved amplified polymorphic sequences (CAPS) markers, 2B-448 on chromosome 2B and 5B-301 on chromosome 5B, were developed from the representative SNPs of the major common loci Qgc.ahau-2B.3/Qphs.ahau-2B.4 controlling GC/PHS resistance and PHS resistance locus Qphs.ahau-5B.4, respectively. Further validation in 171 Chinese mini-core collections confirmed significant correlations of the two CAPS markers with GC and PHS resistance phenotypes under different environments (P<0.05). Furthermore, the wheat public expression database, transcriptomic sequencing, and gene allelic variation analysis identified TraesCS5B02G545100, which encodes glutaredoxin, as a potential candidate gene linked to Qphs.ahau-5B.4. The new CAPS marker CAPS-356 was then developed based on the SNP (T/C) in the coding sequences (CDS) region of TraesCS5B02G545100. The high-density linkage map of the Jing 411/Hongmangchun 21 recombinant inbred lines (RILs) constructed based on speciﬁc locus ampliﬁed fragment sequencing markers showed that CAPS-356 collocated with a novel QTL for PHS resistance, supporting the role of TraesCS5B02G545100 as the potential candidate gene linked to Qphs.ahau-5B.4. These results provide valuable information for the map-based cloning of Qphs.ahau-5B.4 and breeding of PHS resistant white-grained varieties.

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A single nucleotide substitution in the MATE transporter gene regulates plastochron and many noded dwarf phenotype in barley (Hordeum vulgare L.)

GUO Bao-jian, SUN Hong-wei, QI Jiang, HUANG Xin-yu, HONG Yi, HOU Jian, LÜ Chao, WANG Yu-lin, WANG Fei-fei, ZHU Juan, GUO Gang-gang, XU Ru-gen

2023, 22 (8): 2295-2305. DOI: 10.1016/j.jia.2023.02.006

Abstract （366） PDF in ScienceDirect

In higher plants, the shoot apical meristem produces lateral organs in a regular spacing (phyllotaxy) and timing (plastochron). The molecular analysis of mutants associated with phyllotaxy and plastochron would increase our understanding of the mechanism of shoot architecture formation. In this study, we identified mutant mnd8^ynp5 that shows an increased rate of leaf emergence and a larger number of nodes in combination with a dwarfed growth habit from an EMS-treated population of the elite barley cultivar Yangnongpi 5. Using a map-based cloning strategy, the mnd8 gene was narrowed down to a 6.7-kb genomic interval on the long arm of chromosome 5H. Sequence analysis revealed that a C to T single-nucleotide mutation occurred at the first exon (position 953) of HORVU5Hr1G118820, leading to an alanine (Ala) to valine (Val) substitution at the 318th amino acid site. Next, HORVU5Hr1G118820 was defined as the candidate gene of MND8 encoding 514 amino acids and containing two multidrug and toxic compound extrusion (MATE) domains. It is highly homologous to maize Bige1 and has a conserved function in the regulation of plant development by controlling the leaf initiation rate. Examination of modern barely varieties showed that Hap-1 was the dominant haplotype and was selected in barley breeding around the world. Collectively, our results indicated that mnd8^ynp5 is a novel allele of the HORVU5Hr1G118820 gene that is possibly responsible for the shortened plastochron and many noded dwarf phenotype in barley.

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Identification of genetic loci for grain yield‑related traits in the wheat population Zhongmai 578/Jimai 22

LIU Dan, ZHAO De-hui, ZENG Jian-qi, Rabiu Sani SHAWAI, TONG Jing-yang, LI Ming, LI Fa-ji, ZHOU Shuo, HU Wen-li, XIA Xian-chun, TIAN Yu-bing, ZHU Qian, WANG Chun-ping, WANG De-sen, HE Zhong-hu, LIU Jin-dong, ZHANG Yong

2023, 22 (7): 1985-1999. DOI: 10.1016/j.jia.2022.12.002

Abstract （301） PDF in ScienceDirect

The identification of stable quantitative trait locus (QTL) for yield-related traits and tightly linked molecular markers is important for improving wheat grain yield. In the present study, six yield-related traits in a recombinant inbred line (RIL) population derived from the Zhongmai 578/Jimai 22 cross were phenotyped in five environments. The parents and 262 RILs were genotyped using the wheat 50K single nucleotide polymorphism (SNP) array. A high-density genetic map was constructed with 1 501 non-redundant bin markers, spanning 2 384.95 cM. Fifty-three QTLs for six yield-related traits were mapped on chromosomes 1D (2), 2A (9), 2B (6), 2D, 3A (2), 3B (2), 4A (5), 4D, 5B (8), 5D (2), 7A (7), 7B (3) and 7D (5), which explained 2.7–25.5% of the phenotypic variances. Among the 53 QTLs, 23 were detected in at least three environments, including seven for thousand-kernel weight (TKW), four for kernel length (KL), four for kernel width (KW), three for average grain filling rate (GFR), one for kernel number per spike (KNS) and four for plant height (PH). The stable QTLs QKl.caas-2A.1, QKl.caas-7D, QKw.caas-7D, QGfr.caas-2B.1, QGfr.caas-4A, QGfr.caas-7A and QPh. caas-2A.1 are likely to be new loci. Six QTL-rich regions on 2A, 2B, 4A, 5B, 7A and 7D, showed pleiotropic effects on various yield traits. TaSus2-2B and WAPO-A1 are potential candidate genes for the pleiotropic regions on 2B and 7A, respectively. The pleiotropic QTL on 7D for TKW, KL, KW and PH was verified in a natural population. The results of this study enrich our knowledge of the genetic basis underlying yield-related traits and provide molecular markers for high-yield wheat breeding.

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Effect of high-molecular-weight glutenin subunit Dy10 on wheat dough properties and end-use quality

WANG Yan, GUO Zhen-ru, CHEN Qing, LI Yang, ZHAO Kan, WAN Yong-fang, Malcolm J. HAWKESFORD, JIANG Yun-feng, KONG Li, PU Zhi-en, DENG Mei, JIANG Qian-tao, LAN Xiu-jin, WANG Ji-rui, CHEN Guo-yue, MA Jian, ZHENG You-liang, WEI Yu-ming, QI Peng-fei

2023, 22 (6): 1609-1617. DOI: 10.1016/j.jia.2022.08.041

Abstract （423） PDF in ScienceDirect

High-molecular-weight glutenin subunits (HMW-GSs) are the most critical grain storage proteins that determine the unique processing qualities of wheat. Although it is a part of the superior HMW-GS pair (Dx5+Dy10), the contribution of the Dy10 subunit to wheat processing quality remains unclear. In this study, we elucidated the effect of Dy10 on wheat processing quality by generating and analyzing a deletion mutant (with the Dy10-null allele), and by elucidating the changes to wheat flour following the incorporation of purified Dy10. The Dy10-null allele was transcribed normally, but the Dy10 subunit was lacking. These findings implied that the Dy10-null allele reduced the glutenin:gliadin ratio and negatively affected dough strength (i.e., Zeleny sedimentation value, gluten index, and dough development and stability times) and the bread-making quality; however, it positively affected the biscuit-making quality. The incorporation of various amounts of purified Dy10 into wheat flour had a detrimental effect on biscuit-making quality. The results of this study demonstrate that the Dy10 subunit is essential for maintaining wheat dough strength. Furthermore, the Dy10-null allele may be exploited by soft wheat breeding programs.

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Development and characterization of a novel common wheat–Mexico Rye T1DL·1RS translocation line with stripe rust and powdery mildew resistance

LI Jiao-jiao, ZHAO Li, LÜ Bo-ya, FU Yu, ZHANG Shu-fa, LIU Shu-hui, YANG Qun-hui, WU Jun, LI Jia-chuang, CHEN Xin-hong

2023, 22 (5): 1291-1307. DOI: 10.1016/j.jia.2022.08.039

Abstract （421） PDF in ScienceDirect

Rye (Secale cereale L., 2n=2x=14, RR) is a significant genetic resource for improving common wheat because of its resistance to multiple diseases and abiotic-stress tolerant traits. The 1RS chromosome from the German cultivated rye variety Petkus is critical in wheat breeding. However, its weakened disease resistance highlights the need to identify new resources. In the present study, a novel derived line called D27 was developed from common wheat and Mexico Rye. Cytological observations characterized the karyotype of D27 as 2n=42=21 II. Genomic in situ hybridization indicated that a pair of whole-arm translocated Mexico Rye chromosomes were inherited typically in the mitotic and meiosis stages of D27. Experiments using fluorescence in situ hybridization (FISH) and gliadin electrophoresis showed that D27 lacked wheat 1DS chromosomes. They were replaced by 1RS chromosomes of Mexico Rye, supported by wheat simple-sequence repeat markers, rye sequence characterized amplified region markers, and wheat 40K SNP array analysis. The wheat 1DS chromosomes could not be detected by molecular markers and wheat SNP array, but the presence of rye 1RS chromosomes was confirmed. Agronomic trait assessments indicated that D27 had a higher tiller number and enhanced stripe rust and powdery mildew resistance. In addition, dough properties analysis showed that replacing 1DS led to higher viscosity and lower dough elasticity in D27, which was beneficial for cake making. In conclusion, the novel cytogenetically stable common wheat–Mexico Rye T1DL·1RS translocation line D27 offers excellent potential as outstanding germplasm in wheat breeding programs focusing on disease resistance and yield improvement. Additionally, it can be valuable for researching the rye 1RS chromosome’s genetic diversity.

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Investigation of Aegilops umbellulata for stripe rust resistance, heading date, and the contents of iron, zinc, and gluten protein

SONG Zhong-ping, ZUO Yuan-yuan, XIANG Qin, LI Wen-jia, LI Jian, LIU Gang, DAI Shou-fen, YAN Ze-hong

2023, 22 (4): 1258-1265. DOI: 10.1016/j.jia.2022.08.014

Abstract （303） PDF in ScienceDirect

Aegilops umbellulata (UU) is a wheat wild relative that has potential use in the genetic improvement of wheat. In this study, 46 Ae. umbellulata accessions were investigated for stripe rust resistance, heading date (HD), and the contents of iron (Fe), zinc (Zn), and seed gluten proteins. Forty-two of the accessions were classified as resistant to stripe rust, while the other four accessions were classified as susceptible to stripe rust in four environments. The average HD of Ae. umbellulata was significantly longer than that of three common wheat cultivars (180.9 d vs. 137.0 d), with the exception of PI226500 (138.9 d). The Ae. umbellulata accessions also showed high variability in Fe (69.74–348.09 mg kg^–1) and Zn (49.83–101.65 mg kg^–1) contents. Three accessions (viz., PI542362, PI542363, and PI554399) showed relatively higher Fe (230.96–348.09 mg kg^–1) and Zn (92.46–101.65 mg kg^–1) contents than the others. The Fe content of Ae. umbellulata was similar to those of Ae. comosa and Ae. markgrafii but higher than those of Ae. tauschii and common wheat. Aegilops umbellulata showed a higher Zn content than Ae. tauschii, Ae. comosa, and common wheat, but a lower content than Ae. markgrafii. Furthermore, Ae. umbellulata had the highest proportion of γ-gliadin among all the species investigated (Ae. umbellulata vs. other species=mean 72.11% vs. 49.37%; range: 55.33–86.99% vs. 29.60–67.91%). These results demonstrated that Ae. umbellulata exhibits great diversity in the investigated traits, so it can provide a potential gene pool for the genetic improvement of these traits in wheat.

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Development and characterization of wheat–Aegilops kotschyi 1U^k(1A) substitution line with positive dough quality parameters

JIANG Yun, WANG De-li, HAO Ming, ZHANG Jie, LIU Deng-cai

2023, 22 (4): 999-1008. DOI: 10.1016/j.jia.2022.08.020

Abstract （239） PDF in ScienceDirect

Exploring novel high molecular weight glutenin subunits (HMW-GSs) from wild related species is a strategy to improve wheat processing quality. The objective of the present investigation was to identify the chromosomes of the wheat-alien introgression line N124, derived from the hybridization between Triticum aestivum with Aegilops kotschyi, and characterize the effects on quality-related traits. Fluorescence in situ hybridization karyotypes showed that N124 is a disomic 1U^k(1A) substitution line. Sodium dodecyl sulfate polyacrylamide gel electrophoresis (SDS-PAGE) and reversed-phase high-performance liquid chromatography verified N124 expressed two HMW-GSs of the Ae. kotschyi parent. PacBio RNA sequencing and phylogenetic analysis confirmed that the two HMW-GSs were U^kx and U^ky. Compared to the wheat parent, the substitution line had no obvious agronomic defects except fewer grains per spike but improved several major quality parameters. It can be served as a donor or bridge material for wheat quality improvement.

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The miR164-TaNAC14 module regulates root development and abiotic-stress tolerance in wheat seedlings

CHI Qing, DU Lin-ying, MA Wen, NIU Ruo-yu, WU Bao-wei, GUO Li-jian, MA Meng, LIU Xiang-li, ZHAO Hui-xian

2023, 22 (4): 981-998. DOI: 10.1016/j.jia.2022.08.016

Abstract （497） PDF in ScienceDirect

Previous studies have revealed the miR164 family and the miR164-targeted NAC transcription factor genes in rice (Oryza sativa) and Arabidopsis that play versatile roles in developmental processes and stress responses. In wheat (Triticum aestivum L.), we found nine genetic loci of tae-miR164 (tae-MIR164 a to i) producing two mature sequences that down-regulate the expression of three newly identified target genes of TaNACs (TaNAC1, TaNAC11, and TaNAC14) by the cleavage of the respective mRNAs. Overexpression of tae-miR164 or one of its target genes (TaNAC14) demonstrated that the miR164-TaNAC14 module greatly affects root growth and development and stress (drought and salinity) tolerance in wheat seedlings, and TaNAC14 promotes root growth and development in wheat seedlings and enhances drought tolerance, while tae-miR164 inhibits root development and reduces drought and salinity tolerance by down-regulating the expression of TaNAC14. These findings identify the miR164-TaNAC14 module as well as other tae-miR164-regulated genes which can serve as new genetic resources for stress-resistance wheat breeding.

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Advances in studies on the physiological and molecular regulation of barley tillering

Asad RIAZ, Ahmad M. ALQUDAH, Farah KANWAL, Klaus PILLEN, YE Ling-zhen, DAI Fei, ZHANG Guo-ping

2023, 22 (1): 1-13. DOI: 10.1016/j.jia.2022.08.011

Abstract （325） PDF in ScienceDirect

Tillering is a crucial trait closely associated with yield potential and environmental adaptation in cereal crops, regulated by the synergy of endogenous (genetic) and exogenous (environmental) factors. The physiological and molecular regulation of tillering has been intensively studied in rice and wheat. However, tillering research on barley is scarce. This review used the recent advances in bioinformatics to map all known and potential barley tiller development genes with their chromosomal genetic and physical positions. Many of them were mapped for the first time. We also discussed tillering regulation at genetic, physiological, and environmental levels. Moreover, we established a novel link between the genetic control of phytohormones and sugars with tillering. We provided evidence of how environmental cues and cropping systems help optimize the tiller number. This comprehensive review enhances the understanding of barley’s physiological and genetic mechanisms controlling tillering and other developmental traits.

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Analysis of combining ability for stem-related traits and its correlations with lodging resistance heterosis in hybrid wheat

YANG Wei-bing, QIN Zhi-lie, SUN Hui, HOU Qi-ling, GAO Jian-gang, CHEN Xian-chao, ZHANG Li-ping, WANG Yong-bo, ZHAO Chang-ping, ZHANG Feng-ting

2022, 21 (1): 26-35. DOI: 10.1016/S2095-3119(20)63408-6

Abstract （243） PDF in ScienceDirect

With the application of hybrid wheat, lodging is becoming one of the major factors limiting high yield in its production. However, few studies have focused on combining ability and heterosis analysis of stem-related traits. In this study, 24 crosses were made according to NCII genetic design, using the three (photo-sensitive male sterile lines)×eight (restorer lines) incomplete diallel crosses. The length of basal second internode (LBSI) and breaking strength of basal second internode (BSBSI) as well as other stem-related traits were used to perform the principal component analysis (PCA), combining ability and heterosis analysis. The PCA results showed that the variables could be classified into two main factors, which were named as the positive factor (factor 1) and the negative factor (factor 2), and accounted for 52.3 and 33.2%, respectively, of the total variance in different variables, combined with the analysis for index weight indicated that the factor 1-related traits play positive roles in lodging resistance formation of hybrids. Combining ability variance analysis indicated that its genetic performance was mainly dominated by additive gene effects, and the hybrid combinations with higher lodging resistance can be selected by using of 14GF6085 (R1), 14GF6343-2 (R4), 14GF6937 (R6), 14GF7433-1 (R7), and BS1086 (M3), which are with the features with lower general combining ability (GCA) effects of factor 2-related traits whereas higher GCA effects of factor 1-related traits. The heterosis analysis showed that the wide range of heterosis varied with the traits and combinations, and GCA or specific combining ability (SCA) effects of factor 1-related traits except wall thickness of basal second internode (WTBSI) were positively and closely related to the heterosis of lodging resistance. Generally, the correlation coefficients of heterosis to GCA effects of sterile lines (GCA_m) of factor 1-related traits are significantly higher than that to GCA of restorer lines (GCA_r) and SCA, combined with the higher GCA_m variance values of factor 1-related traits compared to GCA_r, the GCA_m of factor 1-related traits should be particularly considered when breeding hybrid combinations. The heritability analysis showed that the narrow-sense heritability of the diameter of basal second internode (DBSI) and the center of gravity height (TCGH) were obviously lower (<60%) than other traits, suggesting that these two traits were suitable for selection in higher generation for parental breeding. These could provide a theoretical basis for parental breeding and heterosis utilization of lodging resistance.

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Characterization of transgenic wheat lines expressing maize ABP7 involved in kernel development

Zaid CHACHAR, Siffat Ullah KHAN, ZHANG Xue-huan, LENG Peng-fei, ZONG Na, ZHAO Jun

2023, 22 (2): 389-399. DOI: 10.1016/j.jia.2022.08.052

Abstract （200） PDF in ScienceDirect

Wheat is one of the major food crops in the world. Functional validation of the genes in increasing the grain yield of wheat by genetic engineering is essential for feeding the ever-growing global population. This study investigated the role of ABP7, a bHLH transcription factor from maize involved in kernel development, in regulating grain yield-related traits in transgenic wheat. Molecular characterization showed that transgenic lines HB123 and HB287 contained multicopy integration of ABP7 in the genome with higher transgene expression. At the same time, QB205 was a transgenic event of single copy insertion with no significant difference in ABP7 expression compared to wild-type (WT) plants. Phenotyping under field conditions showed that ABP7 over-expressing transgenic lines HB123 and HB287 exhibited improved grain yield-related traits (e.g., grain number per spike, grain weight per spike, thousand-grain weight, grain length, and grain width) and increased grain yield per plot, compared to WT plants, whereas line QB205 did not. In addition, total chlorophyll, chlorophyll a, chlorophyll b, and total soluble sugars were largely increased in the flag leaves of both HB123 and HB287 transgenic lines compared to the WT. These results strongly suggest that ABP7 positively regulates yield-related traits and plot grain yield in transgenic wheat. Consequently, ABP7 can be utilized in wheat breeding for grain yield improvement

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TaABI19 positively regulates grain development in wheat

LIU Yun-chuan, WANG Xiao-lu, HAO Chen-yang, IRSHAD Ahsan, LI Tian, LIU Hong-xia, HOU Jian, ZHANG Xue-yong

2023, 22 (1): 41-51. DOI: 10.1016/j.jia.2022.08.049

Abstract （274） PDF in ScienceDirect

Starch is the most important component in endosperm, and its synthesis is regulated by multiple transcription factors (TFs) in cereals. However, whether the functions of these TFs are conserved or not among cereals unclear yet. Here, we cloned a B3 family TF, named as TaABI19 based on its orthologous in maize (Zea mays L.). Alignment of DNA and protein showed that ABI19 was conserved in maize and wheat (Triticum aestivum L.). We found that TaABI19 was highly expressed in young spike and developing grains and encoded a nucleus-localized transcriptional activator in wheat. The taabi19-b1 null mutants obtained by EMS performed down-regulation of starch synthesis, shorter grain length and lower thousand grain weight (TGW). Furthermore, we provided TaABI19 could bind to the promoters of TaPBF homology genes and enhance their expression. Haplotype association showed that TaABI19-B1 was significantly associated with TGW. We found that Hap2 and Hap3 were favored and underwent positive selection in China wheat breeding. Less than fifty percent in the modern cultivars conveying favored haplotypes indicates TaABI19 still can be considered as target loci for marker-assisted selection breeding to increase TGW in China.

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Identification of genetic locus with resistance to take-all in the wheat-Psathyrostachys huashanica Keng introgression line H148

BAI Sheng-sheng, ZHANG Han-bing, HAN Jing, WU Jian-hui, LI Jia-chuang, GENG Xing-xia, LÜ Bo-ya, XIE Song-feng, HAN De-jun, ZHAO Ji-xin, YANG Qun-hui, WU Jun, CHEN Xin-hong

2021, 20 (12): 3101-3113. DOI: 10.1016/S2095-3119(20)63340-8

Abstract （265） PDF in ScienceDirect

Take-all is a devastating soil-borne disease of wheat (Triticum aestivum L.). Cultivating resistant line is an important measure to control this disease. Psathyrostachys huashanica Keng is a valuable germplasm resource with high resistance to take-all. This study reported on a wheat-P. huashanica introgression line H148 with improved take-all resistance compared with its susceptible parent 7182. To elucidate the genetic mechanism of resistance in H148, the F₂ genetic segregating population of H148×XN585 was constructed. The mixed genetic model analysis showed that the take-all resistance was controlled by two major genes with additive, dominant and epistasis effects. Bulked segregant analysis combined with wheat axiom 660K genotyping array analysis showed the polymorphic SNPs with take-all resistance from P. huashanica alien introgression were mainly distributed on the chromosome 2A. Genotyping of the F₂ population using the KASP marker mapped a major QTL in an interval of 68.8–70.1 Mb on 2AS. Sixty-two genes were found in the target interval of the Chinese Spring reference genome sequence. According to the functional annotation of genes, two protein genes that can improve the systematic resistance of plant roots were predicted as candidate genes. The development of wheat-P. huashanica introgression line H148 and the resistant QTL mapping information are expected to provide some valuable references for the fine mapping of disease-resistance gene and development of take-all resistant varieties through molecular marker-assisted selection.

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Genetic effects of Agropyron cristatum 2P chromosome translocation fragments in wheat background

XU Shi-rui, JIANG Bo, HAN Hai-ming, JI Xia-jie, ZHANG Jin-peng, ZHOU Sheng-hui, YANG Xin-ming, LI Xiu-quan, LI Li-hui, LIU Wei-hua

2023, 22 (1): 52-62. DOI: 10.1016/j.jia.2022.08.094

Abstract （196） PDF in ScienceDirect

Agropyron cristatum (2n=4x=28, PPPP) is a wild relative of common wheat which contains a large number of desirable genes that can be exploited for wheat improvement. Wheat–A. cristatum 2P alien translocation lines exhibit many desirable traits, such as small flag leaves, a high spikelet number and density, and a compact plant type. An agronomic trait evaluation and a genetic analysis were carried out on translocation lines and backcross populations of these lines carrying different translocation fragments. The results showed that a translocation fragment from 2PT-3 (2PL) reduced the length of the flag leaves, while translocation fragments from 2PT-3 (2PL) and 2PT-5 (2PL (0.60–1.00)) reduced the width of the flag leaves. A translocation fragment from 2PT-13 (2PS (0.18–0.36)) increased the length and area of the flag leaves. Translocation fragments from 2PT-3 (2PL) and 2PT-8 (2PL (0.86–1.00)) increased the density of spikelets. Translocation fragments from 2PT-7 (2PL (0.00–0.09)), 2PT-8 (2PL (0.86–1.00)), 2PT-10 (2PS), and 2PT-13 (2PS (0.18–0.36)) reduced plant height. This study provides a scientific basis for the effective utilization of wheat–A. cristatum translocation lines.

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Genetic dissection of wheat uppermost-internode diameter and its association with agronomic traits in five recombinant inbred line populations at various field environments

LIU Hang, TANG Hua-ping, LUO Wei, MU Yang, JIANG Qian-tao, LIU Ya-xi, CHEN Guo-yue, WANG Ji-rui, ZHENG Zhi, QI Peng-fei, JIANG Yun-feng, CUI Fa, SONG Yin-ming, YAN Gui-jun, WEI Yuming, LAN Xiu-jin, ZHENG You-liang, MA Jian

2021, 20 (11): 2849-2861. DOI: 10.1016/S2095-3119(20)63412-8

Abstract （207） PDF in ScienceDirect

Uppermost-internode diameter (UID) is a key morphological trait associated with spike development and yield potential in wheat. Our understanding of its genetic basis remains largely unknown. Here, quantitative trait loci (QTLs) for UID with high-density genetic maps were identified in five wheat recombinant inbred line (RIL) populations. In total, 25 QTLs for UID were detected in five RIL populations, and they were located on chromosomes 1A, 1D (3 QTL), 2B (2), 2D (3), 3B, 3D, 4A, 4B (3), 4D, 5A (5), 5B (2), 6B, and 7D. Of them, five major and stable QTLs (QUid.sau-2CN-1D.1, QUid.sau-2SY-1D, QUid.sau-QZ-2D, QUid.sau-SC-3D, and QUid.sau-AS-4B) were identified from each of the five RIL populations in multiple environments. QUid.sau-2CN-1D.1, QUid.sau-2SY-1D and QUid.sau-SC-3D are novel QTLs. Kompetitive Allele Specific PCR (KASP) markers tightly linked to them were further investigated for developing near-isogenic lines (NILs) carrying the major loci. Furthermore, candidate genes at these intervals harboring major and stable QTLs were predicted, and they were associated with plant development and water transportation in most cases. Comparison of physical locations of the identified QTL on the ‘Chinese Spring’ reference genome showed that several QTLs including two major ones, QUid.sau-2CN-1D.1 and QUid.sau-2SY-1D, are likely allelic confirming their validity and effectiveness. The significant relationships detected between UID and other agronomic traits and a proper UID were discussed. Collectively, our results dissected the underlying genetic basis for UID in wheat and laid a foundation for further fine mapping and map-based cloning of these QTLs.

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Physiological response of flag leaf and yield formation of winter wheat under different spring restrictive irrigation regimes in the Haihe Plain, China

LIU Xue-jing, YIN Bao-zhong, HU Zhao-hui, BAO Xiao-yuan, WANG Yan-dong, ZHEN Wen-chao

2021, 20 (9): 2343-2359. DOI: 10.1016/S2095-3119(20)63352-4

Abstract （158） PDF in ScienceDirect

In order to identify the optimum period of spring water-restrictive irrigation for winter wheat (Triticum aestivum L.) in the Haihe Plain, China and elucidate its effects on flag leaf senescence and yield formation, field experiments were conducted at the Xinji Experimental Station of Hebei Agricultural University from 2016 to 2019 by using different irrigation regimes in spring, including the conventional regime involving two irrigation periods (control (CK), the 3-leaf unfolding stage and the anthesis stage) and a series of single, restrictive irrigation regimes (SRI) comprising irrigation at the 3-leaf unfolding stage (3LI), 4LI, 5LI, and 6LI. There are five major findings: (1) The senescence (determined by the green leaf area, GLA) in the 4LI treatment occurred moderately earlier than that in CK, showed no significant difference with that in 5LI and 6LI, and occurred significantly later than that in 3LI. (2) Compared with other SRI treatments, the GLA value and photosynthetic rate in 4LI were 14.82 and 20.1% higher, respectively. Microstructural analysis of flag leaf also revealed that the mesophyll cells and chloroplasts were irregularly arranged under drought stress in 3LI and 6LI; however, drought stress had minimal negative effects on the microstructure in 4LI and 5LI. (3) Postponed irrigation in spring could significantly increase superoxide dismutase (SOD) and catalase (CAT) activities in the early stage of grain filling; however, these activities would subsequently decrease. Among the four SRI treatments, the overall enzyme activities were the highest in 4LI, and the combined malondialdehyde (MDA) content in flag leaves in 4LI and 5LI was 14.5% lower on average than that in 3LI and 6LI. (4) The soluble sugar (SS) and proline (Pro) contents in 4LI were the highest among the four SRI treatments; however, they were lower than those in CK. The abscisic acid (ABA) hormone content in 4LI and 5LI was lower than that in 3LI and 6LI, respectively, suggesting a smaller drought stress effect in 4LI and 5LI. (5) In two growing seasons, there was a larger number of spikes per unit area in 4LI (i.e., 13.4% higher than that in 5LI and 6LI) and the 1 000-grain weight in 4LI was the highest among the four SRI treatments (i.e., 6.0% higher than that in the other three SRI treatments). Therefore, a single restrictive irrigation regime at the 4-leaf unfolding stage is recommended to be effective in slowing down the senescence process of flag leaves and achieving high yield.

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Genetic dissection of the grain filling rate and related traits through linkage analysis and genome-wide association study in bread wheat

YU Hai-xia, DUAN Xi-xian, SUN Ai-qing, SUN Xiao-xiao, ZHANG Jing-juan, SUN Hua-qing, SUN Yan-yan, NING Tang-yuan, TIAN Ji-chun, WANG Dong-xue, LI Hao, FAN Ke-xin, WANG Ai-ping, MA Wu-jun, CHEN Jian-sheng

2022, 21 (10): 2805-2817. DOI: 10.1016/j.jia.2022.07.032

Abstract （215） PDF in ScienceDirect

Wheat grain yield is generally sink-limited during grain filling. The grain-filling rate (GFR) plays a vital role but is poorly studied due to the difficulty of phenotype surveys. This study explored the grain-filling traits in a recombinant inbred population and wheat collection using two highly saturated genetic maps for linkage analysis and genome-wide association study (GWAS). Seventeen stable additive quantitative trait loci (QTLs) were identified on chromosomes 1B, 4B, and 5A. The linkage interval between IWB19555 and IWB56078 showed pleiotropic effects on GFR₁, GFR_max, kernel length (KL), kernel width (KW), kernel thickness (KT), and thousand kernel weight (TKW), with the phenotypic variation explained (PVE) ranging from 13.38% (KW) to 33.69% (TKW). 198 significant marker-trait associations (MTAs) were distributed across most chromosomes except for 3D and 4D. The major associated sites for GFR included IWB44469 (11.27%), IWB8156 (12.56%) and IWB24812 (14.46%). Linkage analysis suggested that IWB35850, identified through GWAS, was located in approximately the same region as QGFRmax2B.3-11, where two high-confidence candidate genes were present. Two important grain weight (GW)-related QTLs colocalized with grain-filling QTLs. The findings contribute to understanding the genetic architecture of the GFR and provide a basic approach to predict candidate genes for grain yield trait QTLs.

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Identification and validation of novel loci associated with wheat quality through a genome-wide association study

PU Zhi-en, YE Xue-ling, LI Yang, SHI Bing-xin, GUO Zhu, DAI Shou-fen, MA Jian, LIU Ze-hou, JIANG Yun-feng, LI Wei, JIANG Qian-tao, CHEN Guo-yue, WEI Yu-ming, ZHENG You-liang

2022, 21 (11): 3131-3147. DOI: 10.1016/j.jia.2022.08.085

Abstract （203） PDF in ScienceDirect

Understanding the genetic basis of quality-related traits contributes to the improvement of grain protein concentration (GPC), grain starch concentration (GSC), and wet gluten concentration (WGC) in wheat, a genome-wide association study (GWAS) based on a mixed linear model (MLM) was performed on the 236 wheat accessions including 160 cultivars and 76 landraces using 55K single nucleotide polymorphism (SNP) array in multiple environments. A total of twelve stable QTL/SNPs were identified to control different quality traits in this populations at least two environments under stripe rust stress; three, seven and two QTLs associated with GPC, GSC, and WGC were characterized respectively and located on chromosomes 1B, 1D, 2A, 2B, 2D, 3B, 3D, 5D, and 7D with the range of phenotypic variation explained (PVE) from 4.2 to 10.7%. Compared with the previously reported QTLs/genes, five QTLs (QGsc.sicau-1BL, QGsc.sicau-1DS, QGsc.sicau-2DL.1, QGsc.sicau-2DL.2, QWgc.sicau-5DL) were potentially novel. KASP markers for SNPs AX-108770574 and AX-108791420 on chromosome on 5D associated with wet gluten concentration were successfully developed. Phenotype of the cultivars containing the A-allele in AX-108770574 and T-allele in AX-108791420 were extremely significantly (P<0.01) higher than that of the landraces containing the G-allele or C-allele of wet gluten concentration in each of the environments. The developed and validated KASP markers could be utilized in molecular breeding aiming to improve the quality in wheat.

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Heredity and gene mapping of a novel white stripe leaf mutant in wheat

LI Hui-juan, JIAO Zhi-xin, NI Yong-jing, JIANG Yu-mei, LI Jun-chang, PAN Chao, ZHANG Jing, SUN Yu-long, AN Jun-hang, LIU Hong-jie, LI Qiao-yun, NIU Ji-shan

2021, 20 (7): 1743-1752. DOI: 10.1016/S2095-3119(20)63345-7

Abstract （133） PDF in ScienceDirect

Spotted leaf (spl) mutant is a type of leaf lesion mimic mutants in plants. We obtained some lesion mimic mutants from ethyl methane sulfonate (EMS)-mutagenized wheat (Triticum aestivum L.) cultivar Guomai 301 (wild type, WT), and one of them was named as white stripe leaf (wsl) mutant because of the white stripes on its leaves. Here we report the heredity and gene mapping of this novel wheat mutant wsl. There are many small scattered white stripes on the leaves of wsl throughout its whole growth period. As the plants grew, the white stripes became more severe and the necrotic area expanded. The mutant wsl grew only weakly before the jointing stage and gradually recovered after jointing. The length and width of the flag leaf, spike number per plant and thousand-grain weight of wsl were significantly lower than those of the WT. Genetic analysis indicated that the trait of white stripe leaf was controlled by a recessive gene locus, named as wsl, which was mapped on the short arm of chromosome 6B by SSR marker assay. Four SSR markers in the F₂population of wsl×CS were linked to wsl in the order of Xgpw1079–Xwmc104–Xgwm508-wsl–Xgpw7651 at 7.1, 5.2, 8.7, and 4.4 cM, respectively and three SSR markers in the F₂ population of wsl×Jimai 22 were linked to wsl in the order of Xgwm508–Xwmc494–Xgwm518-wsl at 3.5, 1.6 and 8.2 cM, respectively. In comparison to the reference genome sequence of Chinese Spring (CS), wsl is located in a 91-Mb region from 88 Mb (Xgwm518) to 179 Mb (Xgpw7651) on chromosome 6BS. Mutant wsl is a novel germplasm for studying the molecular mechanism of wheat leaf development.

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QTL mapping of seedling biomass and root traits under different nitrogen conditions in bread wheat (Triticum aestivum L.)

YANG Meng-jiao, WANG Cai-rong, Muhammad Adeel HASSAN, WU Yu-ying, XIA Xian-chun, SHI Shu-bing, XIAO Yong-gui, HE Zhong-hu

2021, 20 (5): 1180-1192. DOI: 10.1016/S2095-3119(20)63192-6

Abstract （184） PDF in ScienceDirect

Plant nitrogen assimilation and use efficiency in the seedling’s root system are beneficial for adult plants in field condition for yield enhancement. Identification of the genetic basis between root traits and N uptake plays a crucial role in wheat breeding. In the present study, 198 doubled haploid lines from the cross of Yangmai 16/Zhongmai 895 were used to identify quantitative trait loci (QTLs) underpinning four seedling biomass traits and five root system architecture (RSA) related traits. The plants were grown under hydroponic conditions with control, low and high N treatments (Ca(NO₃)₂·4H₂O at 0, 0.05 and 2.0 mmol L⁻¹, respectively). Significant variations among the treatments and genotypes, and positive correlations between seedling biomass and RSA traits (r=0.20 to 0.98) were observed. Inclusive composite interval mapping based on a high-density map from the Wheat 660K single nucleotide polymorphisms (SNP) array identified 51 QTLs from the three N treatments. Twelve new QTLs detected on chromosomes 1AL (1) in the control, 1DS (2) in high N treatment, 4BL (5) in low and high N treatments, and 7DS (3) and 7DL (1) in low N treatments, are first reported in influencing the root and biomass related traits for N uptake. The most stable QTLs (RRS.caas-4DS) on chromosome 4DS, which were related to ratio of root to shoot dry weight trait, was in close proximity of the Rht-D1 gene, and it showed high phenotypic effects, explaining 13.1% of the phenotypic variance. Twenty-eight QTLs were clustered in 12 genetic regions. SNP markers tightly linked to two important QTLs clusters C10 and C11 on chromosomes 6BL and 7BL were converted to kompetitive allele-specific PCR (KASP) assays that underpin important traits in root development, including root dry weight, root surface area and shoot dry weight. These QTLs, clusters and KASP assays can greatly improve the efficiency of selection for root traits in wheat breeding programmes.

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Identification of QTL and underlying genes for root system architecture associated with nitrate nutrition in hexaploid wheat

Marcus GRIFFITHS, Jonathan A. ATKINSON, Laura-Jayne GARDINER, Ranjan SWARUP, Michael P. POUND, Michael H. WILSON, Malcolm J. BENNETT, Darren M. WELLS

2022, 21 (4): 917-932. DOI: 10.1016/S2095-3119(21)63700-0

Abstract （185） PDF in ScienceDirect

The root system architecture (RSA) of a crop has a profound effect on the uptake of nutrients and consequently the potential yield. However, little is known about the genetic basis of RSA and resource adaptive responses in wheat (Triticum aestivum L.). Here, a high-throughput germination paper-based plant phenotyping system was used to identify seedling traits in a wheat doubled haploid mapping population, Savannah×Rialto. Significant genotypic and nitrate-N treatment variation was found across the population for seedling traits with distinct trait grouping for root size-related traits and root distribution-related traits. Quantitative trait locus (QTL) analysis identified a total of 59 seedling trait QTLs. Across two nitrate treatments, 27 root QTLs were specific to the nitrate treatment. Transcriptomic analyses for one of the QTLs on chromosome 2D, which was found under low nitrate conditions, revealed gene enrichment in N-related biological processes and 28 differentially expressed genes with possible involvement in a root angle response. Together, these findings provide genetic insight into root system architecture and plant adaptive responses to nitrate, as well as targets that could help improve N capture in wheat.

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Allele mining of wheat ABA receptor at TaPYL4 suggests neo-functionalization among the wheat homoeologs

WU Bang-bang, SHI Meng-meng, Mohammad POURKHEIRANDISH, ZHAO Qi, WANG Ying, YANG Chen-kang, QIAO Ling, ZHAO Jia-jia, YAN Su-xian, ZHENG Xing-wei, ZHENG Jun

2022, 21 (8): 2183-2196. DOI: 10.1016/S2095-3119(21)63699-7

Abstract （213） PDF in ScienceDirect

ABA receptors (PYR/PYL/RCAR) play a central role in the water loss control of plants. A previous report indicated that TaPYL4 is a critical gene in wheat that improves grain production under drought conditions and increases water use efficiency. In this study, we analyzed the sequence polymorphisms and genetic effects of TaPYL4s. Based on isolated TaPYL4 genes from chromosomes 2A, 2B and 2D, three haplotypes were detected in the promoter region of TaPYL4-2A, and two haplotypes were present in TaPYL4-2B and TaPYL4-2D, respectively. Marker/trait association analysis indicated that TaPYL4-2A was significantly associated with plant height in 262 Chinese wheat core collection accessions, as well as the drought tolerance coefficient (DTC) for plant height in 239 wheat varieties from Shanxi Province in multiple environments. However, the frequencies of favored drought-tolerant haplotype TaPYL4-2A-Hap2 were considerably low, accounting for only 10%, and lines with this certain Hap could be reserved in the breeding program. TaPYL4-2B was significantly associated with grain number, and the favored haplotype TaPYL4-2B-Hap1 was the dominant allele of above 90% in the collection. For TaPYL4-2D, there were no significant differences in these traits between the two haplotypes in either of the two panels. These results indicate that variation might lead to functional differentiation among the homoeologs and the haplotypes had undergone artificial selection during breeding. Two molecular markers developed to distinguish these haplotypes could be used for breeding in water-limited regions.

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Variations in the quality parameters and gluten proteins in synthetic hexaploid wheats solely expressing the Glu-D1 locus

DAI Shou-fen, CHEN Hai-xia, LI Hao-yuan, YANG Wan-jun, ZHAI Zhi, LIU Qian-yu, LI Jian, YAN Ze-hong

2022, 21 (7): 1877-1885. DOI: 10.1016/S2095-3119(21)63651-1

Abstract （196） PDF in ScienceDirect

This study evaluated the quality potential of seven synthetic hexaploid wheats (2n=6x=42, AABBDD) expressing only allelic variation at Glu-D1 of Aegilops tauschii (SHWSD). Major quality parameters related to dough strength, gluten proteins (including high-molecular-weight glutenin subunits (HMW-GS) and low-molecular-weight glutenin subunits (LMW-GS), gliadins), and their ratios between SHWSD and the weak gluten wheat control Chuannong 16 (CN16) were measured in at least three environments (except STD7). The zeleny sedimentation value (ZSV), dough development time (DDT), dough stability time (DST), and farinograph quality number (FQN) of SHWSD were considered stable under different environments, with their respective ranges being 8.00–17.67 mL, 0.57–1.50 min, 0.73–1.80 min, and 9.50–27.00. The ZSV, DDT, DST, and FQN of SHWSD were smaller than those of CN16, suggesting that SHWSD had a weaker dough strength than CN16. Although SHWSD had a lower gluten index than CN16, its wet and dry gluten contents were similar to or even higher than those of CN16 in all environments tested. The protein content of grains (12.81–18.21%) and flours (14.20–20.31%) in SHWSD was higher than that in CN16. The amount of HMW-GS in SHWSD sharply decreased under the expression of fewer HMW-GS genes, and the LMW-GS, gliadins, and total glutenins were simultaneously increased in SHWSD in comparison with CN16. Moreover, SHWSD had higher ratios of LMW-GS/glutenin and gliadin/glutenin but a lower ratio of HMW-GS/glutenin than CN16. These results provide necessary information for the utilization of SHWSD in weak-gluten wheat breeding.

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Influence of high-molecular-weight glutenin subunit deletions at the Glu-A1 and Glu-D1 loci on protein body development, protein components and dough properties of wheat (Triticum aestivum L.)

LIU Da-tong, ZHANG Xiao, JIANG Wei, LI Man, WU Xu-jiang, GAO De-rong, BIE Tong-de, LU Cheng-bin

2022, 21 (7): 1867-1876. DOI: 10.1016/S2095-3119(21)63605-5

Abstract （264） PDF in ScienceDirect

High-molecular-weight glutenin subunits (HMW-GSs) play a critical role in determining the viscoelastic properties of wheat. As the organelle where proteins are stored, the development of protein bodies (PBs) reflects the status of protein synthesis and also affects grain quality to a great extent. In this study, with special materials of four near-isogenic lines in a Yangmai 18 background we created, the effects of Glu-A1 and Glu-D1 loci deletions on the development and morphological properties of the protein body, protein components and dough properties were investigated. The results showed that the deletion of the HMW-GS subunit delayed the development process of the PBs, and slowed the increases of volume and area of PBs from 10 days after anthesis (DAA) onwards. In contrast, the areas of PBs at 25 DAA, the middle or late stage of endosperm development, showed no distinguishable differences among the four lines. Compared to the wild type and single null type in Glu-A1, the ratios of HMW-GSs to low-molecular-weight glutenin subunits (LMW-GSs), glutenin macropolymer (GMP) content, mixograph parameters as well as extension parameters decreased in the single null type in Glu-D1 and double null type in Glu-A1 and Glu-D1, while the ratios of gliadins (Gli)/glutenins (Glu) in those types increased. The absence of Glu-D1 subunits decreased both dough strength and extensibility significantly compared to the Glu-A1 deletion type. These results provide a detailed description of the effect of HMW-GS deletion on PBs, protein traits and dough properties, and contribute to the utilization of Glu-D1 deletion germplasm in weak gluten wheat improvement for use in cookies, cakes and southern steamed bread in China and liquor processing.

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A major and stable QTL for wheat spikelet number per spike validated in different genetic backgrounds

DING Pu-yang, MO Zi-qiang, TANG Hua-ping, MU Yang, DENG Mei, JIANG Qian-tao, LIU Ya-xi, CHEN Guang-deng, CHEN Guo-yue, WANG Ji-rui, LI Wei, QI Peng-fei, JIANG Yun-feng, KANG Hou-yang, YAN Gui-jun, Wei Yu-ming, ZHENG You-liang, LAN Xiu-jin, MA Jian

2022, 21 (6): 1551-1562. DOI: 10.1016/S2095-3119(20)63602-4

Abstract （267） PDF in ScienceDirect

The spikelet number per spike (SNS) contributes greatly to grain yield in wheat. Identifying various genes that control wheat SNS is vital for yield improvement. This study used a recombinant inbred line population genotyped by the Wheat55K single-nucleotide polymorphism array to identify two major and stably expressed quantitative trait loci (QTLs) for SNS. One of them (QSns.sau-2SY-2D.1) was reported previously, while the other (QSns.sau-2SY-7A) was newly detected and further analyzed in this study. QSns.sau-2SY-7A had a high LOD value ranging from 4.46 to 16.00 and explained 10.21–40.78% of the phenotypic variances. QSns.sau-2SY-7A was flanked by the markers AX-110518554 and AX-110094527 in a 4.75-cM interval on chromosome arm 7AL. The contributions and interactions of both major QTLs were further analyzed and discussed. The effect of QSns.sau-2SY-7A was successfully validated by developing a tightly linked kompetitive allele specific PCR marker in an F_2:3 population and a panel of 101 high-generation breeding wheat lines. Furthermore, several genes including the previously reported WHEAT ORTHOLOG OF APO1 (WAPO1), an ortholog of the rice gene ABERRANT PANICLE ORGANIZATION 1 (APO1) related to SNS, were predicted in the interval of QSns.sau-2SY-7A. In summary, these results revealed the genetic basis of the multi-spikelet genotype of wheat line 20828 and will facilitate subsequent fine mapping and breeding utilization of the major QTLs.

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TaSnRK2.4 is a vital regulator in control of thousand-kernel weight and response to abiotic stress in wheat

MIAO Li-li, LI Yu-ying, ZHANG Hong-juan, ZHANG Hong-ji, LIU Xiu-lin, WANG Jing-yi, CHANG Xiao-ping, MAO Xin-guo, JING Rui-lian

2021, 20 (1): 46-54. DOI: 10.1016/S2095-3119(19)62830-3

Abstract （173） PDF in ScienceDirect

Sucrose non-fermenting 1-related protein kinase 2 (SnRK2) is a plant-specific serine/threonine kinase involved in response to adverse environmental stimuli. Previous studies showed that TaSnRK2.4 was involved in response to abiotic stresses and conferred enhanced tolerance to multiple stresses in Arabidopsis. Further experiments were performed to decipher the underlying mechanisms and discover new functions. The genomic sequences of TaSnRK2.4s locating on chromosome 3A, 3B and 3D were obtained. Sequencing identified one and 13 variations of TaSnRK2.4-3A and TaSnRK2.4-3B, respectively, but no variation was detected in TaSnRK2.4-3D. The markers 2.4AM1, 2.4BM1 and 2.4BM2 were developed based on three variations. Association analysis showed that both TaSnRK2.4-3A and TaSnRK2.4-3B were significantly associated with thousand-kernel weight (TKW), and that SNP3A-T and SNP3B-C were favorable alleles for higher TKW. Yeast two-hybrid and split luciferase assays showed that TaSnRK2.4 physically interacted with abiotic stress responsive protein TaLTP3, suggesting that TaSnRK2.4 enhanced abiotic stress tolerance by activating TaLTP3. Our studies suggested that TaSnRK2.4 have potential in improving TKW and response to abiotic stress.

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TaIAA15 genes regulate plant architecture in wheat

LI Fu, YAN Dong, GAO Li-feng, LIU Pan, ZHAO Guang-yao, JIA Ji-zeng, REN Zheng-long

2022, 21 (5): 1243-1252. DOI: 10.1016/S2095-3119(20)63480-3

Abstract （335） PDF in ScienceDirect

Bread wheat (Triticum aestivum L.) is one of the most important staple crops worldwide. The phytohormone auxin plays critical roles in the regulation of plant growth and development. However, only a few auxin-related genes have been genetically demonstrated to be involved in the control of plant architecture in wheat thus far. In this study, we characterized an auxin-related gene in wheat, TaIAA15, and found that its ectopic expression in rice decreased the plant height and increased the leaf angle. Correlation analysis indicated that TaIAA15-3B was associated with plant height (Ph), spike length (SL) and 1 000-grain weight (TGW) in wheat, and Hap-II of TaIAA15-3B was the most favored allele and selected by modern breeding in China. This study sheds light on the role of auxin signaling on wheat plant architecture as well as yield related traits.

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The wheat receptor-like cytoplasmic kinase TaRLCK1B is required for host immune response to the necrotrophic pathogen Rhizoctonia cerealis

WU Tian-ci, ZHU Xiu-liang, LÜ Liang-jie, CHEN Xi-yong, XU Gang-biao, ZHANG Zeng-yan

2020, 19 (11): 2616-2627. DOI: 10.1016/S2095-3119(20)63160-4

Abstract （145） PDF in ScienceDirect

Receptor-like cytoplasmic kinases (RLCKs) represent a large family of proteins in plants. In Arabidopsis and rice, several RLCKs in subfamily VII (RLCKs-VII) have been implicated in pathogen-associated molecular pattern-triggered immunity and basal resistance against bacterial and fungal pathogens. However, little is known about roles of RLCKs-VII of the important crop common wheat (Triticum aestivum) in immune responses. Here, we isolated a RLCK-VII-encoding gene from wheat, designated as TaRLCK1B, and investigated its role in host immune response to infection of a necrotrophic fungus Rhizoctonia cerealis that is a major pathogen of sharp eyespot, a destructive disease of wheat. RNA-sequencing and RT-qPCR analyses showed that transcriptional level of TaRLCK1B was significantly higher in sharp eyespot-resistant wheat cultivars than in susceptible wheat cultivars. The gene transcription was rapidly and markedly elevated in the resistant wheat cultivars by R. cerealis infection. The TaRLCK1B protein was closely related to OsRLCK176, a rice resistance-related RLCKs-VII, with 84.03% identity. Virus-induced gene silencing plus wheat response to R. cerealis assay results indicated that silencing of TaRLCK1 impaired resistance to R. cerealis. Meantime, silencing of TaRLCK1 significantly elevated both the content of H₂O₂ (a major kind of reactive oxygen species, ROS) and the transcriptional level of the ROS-generating enzyme-encoding gene RBOH, but repressed the expression of the ROS-scavenging enzyme-encoding gene CAT1 at 18 hours after inoculation (hai) with R. cerealis. Taken together, these data suggested that TaRLCK1B was required for the early immune response of wheat to R. cerealis through modulating ROS signaling in wheat.

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Grain proteomic analysis reveals central stress responsive proteins caused by wheat-Haynaldia villosa 6VS/6AL translocation

ZOU Rong, WU Ji-su, WANG Ruo-mei, YAN Yue-ming

2020, 19 (11): 2628-2642. DOI: 10.1016/S2095-3119(19)62846-7

Abstract （111） PDF in ScienceDirect

Haynaldia villosa (2n=14, VV), a wild grass of the subtribe Triticeae, serves as potential gene resources for wheat genetic improvement. In this study, the proteome characterization during grain development of Yangmai 5 and Yangmai 5-H. villosa 6VS/6AL translocation line was investigated by a comparative proteomic approach. Two-dimensional electrophoresis identified 81 differentially accumulated proteins (DAPs) during five grain developmental stages in wheat-H. villosa translocation line. These proteins were mainly involved in stress defense, storage protein, energy metabolism, protein metabolism and folding, carbon metabolism, nitrogen metabolism, and starch metabolism. In particular, 6VS/6AL translocation led to significant upregulation of 36 DAPs and specific expression of 11 DAPs such as chitinase, thaumatin-like proteins, glutathione transferase, α-amylase inhibitor, heat shock proteins, and betaine aldehyde dehydrogenase. These proteins mainly involved in biotic and abiotic stress responses. Further analysis found that the upstream 1 500 promoter regions of these stress-responsive DAP genes contained multiple high-frequency cis-acting elements related to stress defense such as abscisic acid response element ABRE, methyl jasmonate (MeJA)-response element TGACG-motif and CGTCA-motif involved in oxidative stress and antioxidant response element (ARE). RNA-seq and RT-qPCR analyses revealed the high expression of these stress-defensive DAP genes in the developing grains, particularly at the early-middle grain filling stages. Our results demonstrated that 6VS chromosome of H. villosa contains abundant stress-defensive proteins that have potential values for wheat genetic improvement.

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Quantitative trait loci analysis for root traits in synthetic hexaploid wheat under drought stress conditions

LIU Rui-xuan, WU Fang-kun, YI Xin, LIN Yu, WANG Zhi-qiang, LIU Shi-hang, DENG Mei, MA Jian, WEI Yu-ming, ZHENG You-liang, LIU Ya-xi

2020, 19 (8): 1947-1960. DOI: 10.1016/S2095-3119(19)62825-X

Abstract （187） PDF in ScienceDirect

Synthetic hexaploid wheat (SHW), possesses numerous genes for drought that can help breeding for drought-tolerant wheat varieties. We evaluated 10 root traits at seedling stage in 111 F₉ recombinant inbred lines derived from a F₂ population of a SHW line (SHW-L1) and a common wheat line, under normal (NC) and polyethylene glycol-simulated drought stress conditions (DC). We mapped quantitative trait loci (QTLs) for root traits using an enriched high-density genetic map containing 120 370 single nucleotide polymorphisms (SNPs), 733 diversity arrays technology markers (DArT) and 119 simple sequence repeats (SSRs). With four replicates per treatment, we identified 19 QTLs for root traits under NC and DC, and 12 of them could be consistently detected with three or four replicates. Two novel QTLs for root fresh weight and root diameter under NC explained 9 and 15.7% of the phenotypic variation respectively, and six novel QTLs for root fresh weight, the ratio of root water loss, total root surface area, number of root tips, and number of root forks under DC explained 8.5–14% of the phenotypic variation. Here seven of eight novel QTLs could be consistently detected with more than three replicates. Results provide essential information for fine-mapping QTLs related to drought tolerance that will facilitate breeding drought-tolerant wheat cultivars.

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Genome-wide identification and transcriptome profiling reveal great expansion of SWEET gene family and their wide-spread responses to abiotic stress in wheat (Triticum aestivum L.)

QIN Jin-xia, JIANG Yu-jie, LU Yun-ze, ZHAO Peng, WU Bing-jin, LI Hong-xia, WANG Yu, XU Sheng-bao, SUN Qi-xin, LIU Zhen-shan

2020, 19 (7): 1704-1720. DOI: 10.1016/S2095-3119(19)62761-9

Abstract （193） PDF in ScienceDirect

The Sugars Will Eventually be Exported Transporter (SWEET) gene family, identified as sugar transporters, has been demonstrated to play key roles in phloem loading, grain filling, pollen nutrition, and plant-pathogen interactions. To date, the study of SWEET genes in response to abiotic stress is very limited. In this study, we performed a genome-wide identification of the SWEET gene family in wheat and examined their expression profiles under mutiple abiotic stresses. We identified a total of 105 wheat SWEET genes, and phylogenic analysis revealed that they fall into five clades, with clade V specific to wheat and its closely related species. Of the 105 wheat SWEET genes, 59% exhibited significant expression changes after stress treatments, including drought, heat, heat combined with drought, and salt stresses, and more up-regulated genes were found in response to drought and salt stresses. Further hierarchical clustering analysis revealed that SWEET genes exhibited differential expression patterns in response to different stress treatments or in different wheat cultivars. Moreover, different phylogenetic clades also showed distinct response to abiotic stress treatments. Finally, we found that homoeologous SWEET genes from different wheat subgenomes exhibited differential expression patterns in response to different abiotic stress treatments. The genome-wide analysis revealed the great expansion of SWEET gene family in wheat and their wide participation in abiotic stress response. The expression partitioning of SWEET homoeologs under abiotic stress conditions may confer greater flexibility for hexaploid wheat to adapt to ever changing environments.

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Genetic analysis and QTL mapping of a novel reduced height gene in common wheat (Triticum aestivum L.)

ZHOU Chun-yun, XIONG Hong-chun, LI Yu-ting, GUO Hui-jun, XIE Yong-dun, ZHAO Lin-shu, GU Jiayu, ZHAO Shi-rong, DING Yu-ping, SONG Xi-yun, LIU Lu-xiang

2020, 19 (7): 1721-1730. DOI: 10.1016/S2095-3119(20)63224-5

Abstract （184） PDF in ScienceDirect

Low stature in wheat is closely associated with lodging resistance, and this impacts harvest index and grain yield. The discovery of novel dwarfing or semi-dwarfing genes can have great significance for dwarf wheat breeding. In this study, we identified an EMS-induced dwarf wheat mutant JE0124 from the elite cultivar Jing411. JE0124 possesses increased stem strength and a 33% reduction in plant height compared with wild type. Gibberellic acid (GA) treatment analysis suggested that JE0124 was GA-sensitive. Analysis of the frequency distribution of plant height in four F₂ populations derived from crosses between JE0124 and the relatively taller varieties Nongda 5181 and WT indicated that the dwarfism phenotype was quantitatively inherited. We used two F₂ populations and 312 individuals from the reciprocal cross of Nongda 5181 and JE0124 to map the quantitative trait locus (QTL) for reduced height to a 0.85-cM interval on chromosome 2DL. The mapping was done by using a combination of 660K SNP array-based bulked segregant analysis (BSA) and genetic linkage analysis, with logarithm of odds (LOD) scores of 5.34 and 5.78, respectively. Additionally, this QTL accounted for 8.27–8.52% of the variation in the phenotype. The dwarf mutant JE0124 and the newly discovered dwarfing gene on chromosome 2DL in this study will enrich genetic resources for dwarf wheat breeding.

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dCAPS markers developed for nitrate transporter genes TaNRT2L12s associating with 1 000-grain weight in wheat

HUANG Jun-fang, LI Long, MAO Xin-guo, WANG Jing-yi, LIU Hui-min, LI Chao-nan, JING Rui-lian

2020, 19 (6): 1543-1553. DOI: 10.1016/S2095-3119(19)62683-3

Abstract （119） PDF in ScienceDirect

Nitrate transporters (NRTs) are regulators of nitrate assimilation and transport. The genome sequences of TaNRT2L12-A, -B and -D were cloned from wheat (Triticum aestivum L.), and polymorphisms were analyzed by sequencing. TaNRT2L12-D in a germplasm population was highly conserved. However, 38 single nucleotide polymorphisms (SNPs) in TaNRT2L12-A coding region and 11 SNPs in TaNRT2L12-B coding region were detected. Two derived cleaved amplified polymorphic sequences (dCAPS) markers A-CSNP1 and A-CSNP2 were developed for TaNRT2L12-A based on SNP-351 and SNP-729, and three haplotypes were identified in the germplasm population. B-CSNP1 and B-CSNP2 were developed for TaNRT2L12-B based on SNP-237 and SNP-1 227, and three haplotypes were detected in the germplasm population. Association analyses between the markers and agronomic traits in 30 environments and phenotypic comparisons revealed that A-CSNP2-A is a superior allele of shorter plant height (PH), length of penultimate internode (LPI) and peduncle length (PL), B-CSNP2-G is a superior allele of higher grain number per spike (GNS). Hap-6B-1 containing both superior alleles B-CSNP1-C and B-CSNP2-A is a superior haplotype of 1 000-grain weight (TGW). Expression analysis showed that TaNRT2L12-B is mainly expressed in the root base and regulated by nitrate. Therefore, TaNRT2L12 may be involved in nitrate transport and signaling to regulate TGW in wheat. The superior alleles and dCAPS markers of TaNRT2L12-A/B are beneficial to genetic improvement and germplasm enhancement with molecular markers-assisted selection.

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Bioinformatic identification and analyses of the non-specific lipid transfer proteins in wheat

FANG Zheng-wu, HE Yi-qin, LIU Yi-ke, JIANG Wen-qiang, SONG Jing-han, WANG Shu-ping, MA Dong-fang, YIN Jun-liang

2020, 19 (5): 1170-1185. DOI: 10.1016/S2095-3119(19)62776-0

Abstract （141） PDF in ScienceDirect

Non-specific lipid transfer proteins (nsLTPs/LTPs) that can transport various phospholipids across the membrane in vitro are widespread in the plant kingdom, and they play important roles in many biological processes that are closely related to plant growth and development. Recently, nsLTPs have been shown to respond to different forms of abiotic stresses. Despite the vital roles of nsLTPs in many plants, little is known about the nsLTPs in wheat. In this study, 330 nsLTP proteins were identified in wheat and they clustered into five types (1, 2, c, d, and g) by phylogenetic analysis with the nsLTPs from maize, Arabidopsis, and rice. The wheat nsLTPs of type d included three subtypes (d1, d2, and d3) and type g included seven subtypes (g1–g7). Genetic structure and motif pattern analyses showed that members of each type had similar structural composition. Moreover, GPI-anchors were found to exist in non-g type members from wheat for the first time. Chromosome mapping revealed that all five types were unevenly and unequally distributed on 21 chromosomes. Furthermore, gene duplication events contributed to the proliferation of the nsLTP genes. Large-scale data mining of RNA-seq data covering multiple growth stages and numerous stress treatments showed that the transcript levels of some of the nsLTP genes could be strongly induced by abiotic stresses, including drought and salinity, indicating their potential roles in mediating the responses of the wheat plants to these abiotic stress conditions. These findings provide comprehensive insights into the nsLTP family members in wheat, and offer candidate nsLTP genes for further studies on their roles in stress resistance and potential for improving wheat breeding programs.

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Molecular detection of the powdery mildew resistance genes in winter wheats DH51302 and Shimai 26

QU Yun-feng, WU Pei-pei, HU Jing-huang, CHEN Yong-xing, SHI Zhan-liang, QIU Dan, LI Ya-hui, ZHANG Hong-jun, ZHOU Yang, YANG Li, LIU Hong-wei, ZHU Tong-quan, LIU Zhi-yong, ZHANG Yan-ming, LI Hong-jie

2020, 19 (4): 931-940. DOI: 10.1016/S2095-3119(19)62644-4

Abstract （122） PDF in ScienceDirect

Resistance to powdery mildew is an important trait of interest in many wheat breeding programs. The information on genes conferring resistance to powdery mildew in wheat cultivars is useful in parental selection. Winter wheat breeding line DH51302 derived from Liangxing 99 and cultivar Shimai 26 derived from Jimai 22 showed identical infection patterns against 13 isolates of Blumeria graminis f. sp. tritici (Bgt) that causes wheat powdery mildew. DH51302 and Shimai 26 were crossed to a powdery mildew susceptible cultivar Zhongzuo 9504 and the F_2:3 families were used in molecular localization of the resistance genes. Fourteen polymorphic markers, which were linked to Pm52 from Liangxing 99, were used to establish the genetic linkage maps for the resistance genes PmDH51302 and PmSM26 in DH51302 and Shimai 26, respectively. These genes were placed in the same genetic interval where Pm52 resides. Analysis of gene-linked molecular markers indicated that PmDH51302 and PmSM26 differed from other powdery mildew resistance genes on chromosome arm 2BL, such as Pm6, Pm33, Pm51, MlZec1, MlAB10, and Pm64. Based on the results of reaction patterns to different Bgt isolates and molecular marker localization, together with the pedigree information, DH51302 and Shimai 26 carried the same gene, Pm52, which confers their resistance to powdery mildew.

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Heterologous expression of the ThIPK2 gene enhances drought resistance of common wheat

ZHANG Shu-juan, LI Yu-lian, SONG Guo-qi, GAO Jie, ZHANG Rong-zhi, LI Wei, CHEN Ming-li, LI Gen-ying

2020, 19 (4): 941-952. DOI: 10.1016/S2095-3119(19)62714-0

Abstract （129） PDF in ScienceDirect

ThIPK2 is an inositol polyphosphate kinase gene cloned from Thellungiella halophila that participates in diverse cellular processes. Drought is a major limiting factor in wheat (Triticum aestivum L.) production. The present study investigated whether the application of the ThIPK2 gene could increase the drought resistance of transgenic wheat. The codon-optimized ThIPK2 gene was transferred into common wheat through Agrobacterium-mediated transformation driven by either a constitutive maize ubiquitin promoter or a stress-inducible rd29A promoter from Arabidopsis. Molecular characterization confirmed the presence of the foreign gene in the transformed plants. The transgenic expression of ThIPK2 in wheat led to significantly improve drought tolerance compared to that observed in control plants. Compared to the wild type (WT) plants, the transgenic plants showed higher seed germination rates, better developed root systems, a higher relative water content (RWC) and total soluble sugar content, and less cell membrane damage under drought stress conditions. The expression profiles showed different expression patterns with the use of different promoters. The codon-optimized ThIPK2 gene is a candidate gene to enhance wheat drought stress tolerance by genetic engineering.

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Identification of QTL for adult plant resistance to stripe rust in bread wheat line C33

LUO Jiang-tao, ZHENG Jian-min, WAN Hong-shen, YANG Wu-yun, LI Shi-zhao, PU Zong-jun

2020, 19 (3): 624-631. DOI: 10.1016/S2095-3119(19)62638-9

Abstract （158） PDF in ScienceDirect

Stripe rust, caused by Puccinia striiformis f. sp. tritici, is a serious disease in bread wheat (Triticum aestivum L.). Identification and use of adult plant resistance (APR) resources are important for stripe rust resistance breeding. Bread wheat line C33 is an exotic germplasm that has shown stable APR to stripe rust for more than 10 years in Sichuan Province of China. Here, 183 recombinant inbred lines (RILs) derived from the cross between C33 and a susceptible line X440 were genotyped with diversity arrays technology (DArT) markers to identify resistance quantitative trait locus (QTL). Field trials were conducted in five years at Chengdu and Xindu of Sichuan Province, using maximum disease severity (MDS) as stripe rust reaction phenotypes. A total of four quantitative trait loci (QTLs) were detected, respectively designed as QYr.saas-3AS, QYr.saas-5AL, QYr.saas-5BL, and QYr.saas-7DS, explaining 4.14–15.21% of the phenotypic variances. QYr.saas-5BL and QYr.saas-7DS were contributed by C33. However, the level for stripe rust resistance contributed by them was not strong as C33, suggesting the presence of other unidentified QTLs in C33. QYr.saas-7DS corresponded to Yr18 and QYr.saas-5BL remains to be formally named. The RIL lines carrying combinations QYr.saas-5AL, QYr.saas-5BL, and QYr.saas-7DS showed comparability resistance with C33. The present study provides resources to pyramid diverse genes into locally adapted elite germplasm to improve the stripe rust resistance of bread wheat.

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Genetic progress in stem lodging resistance of the dominant wheat cultivars adapted to Yellow-Huai River Valleys Winter Wheat Zone in China since 1964

ZHANG Hong-jun, LI Teng, LIU Hong-wei, MAI Chun-yan, YU Guang-jun, LI Hui-li, YU Li-qiang, MENG Ling-zhi, JIAN Da-wei, YANG Li, LI Hong-jie, ZHOU Yang

2020, 19 (2): 438-448. DOI: 10.1016/S2095-3119(19)62627-4

Abstract （144）

Analysis of genetic progress for lodging-related traits provides important information for further improvement of lodging resistance. Forty winter wheat cultivars widely grown in the Yellow-Huai River Valleys Winter Wheat Zone (YHWZ) of China during the period of 1964–2015 were evaluated for several lodging-related traits in three cropping seasons. Plant height, height at center of gravity, length of the basal second internode, and lodging index decreased significantly in this period, and the average annual genetic gains for these traits were –0.50 cm or –0.62%, –0.27 cm or –0.60%, –0.06 cm or –0.63%, and –0.01 or –0.94%, respectively. Different from other traits, stem strength showed a significant increasing trend with the breeding period, and the annual genetic gains were 0.03 N or 0.05%. Correlation analysis showed that lodging index was positively correlated with plant height, height at center of gravity, and length of the basal second internode, but negatively correlated with stem strength. Meanwhile, significantly positive correlations were observed between plant height, height at center of gravity, and length of the basal first and second internodes. By comparison with the wild types, dwarfing genes had significant effects on all lodging-related traits studied except for length of the basal first internode and stem strength. Principle component analysis demonstrated that plant height and stem strength were the most important factors influencing lodging resistance. Clustering analysis based on the first two principle components further indicated the targets of wheat lodging-resistant breeding have changed from reducing plant height to strengthening stem strength over the breeding periods. This study indicates that the increase of stem strength is vital to improve lodging resistance in this region under the high-yielding condition when plant height is in an optimal range.

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Genetic and agronomic traits stability of marker-free transgenic wheat plants generated from Agrobacterium-mediated co-transformation in T₂ and T₃generations

LIU Hui-yun, WANG Ke, WANG Jing, DU Li-pu, PEI Xin-wu, YE Xing-guo

2020, 19 (1): 23-32. DOI: 10.1016/S2095-3119(19)62601-8

Abstract （99） PDF in ScienceDirect

Genetically modified wheat has not been commercially utilized in agriculture largely due to regulatory hurdles associated with traditional transformation methods. Development of marker-free transgenic wheat plants will help to facilitate biosafety evaluation and the eventual environmental release of transgenic wheat varieties. In this study, the marker-free transgenic wheat plants previously obtained by Agrobacterium-mediated co-transformation of double T-DNAs vector were identified by fluorescence in situ hybridization (FISH) in the T₁ generation, and their genetic stability and agronomic traits were analyzed in T₂ and T₃ generations. FISH analysis indicated that the transgene often integrated into a position at the distal region of wheat chromosomes. Furthermore, we show that the GUS transgene was stably inherited in the marker-free transgenic plants in T₁ to T₃ generations. No significant differences in agronomic traits or grain characteristics were observed in T₃generation, with the exception of a small variation in spike length and grains per spike in a few lines. The selection marker of bar gene was not found in the transgenic plants through T₁ to T₃ generations. The results from this investigation lay a solid foundation for the potential application of the marker-free transgenic wheat plants achieved through the co-transformation of double T-DNAs vector by Agrobacterium in agriculture after biosafty evaluation.

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Biotic and abiotic stress-responsive genes are stimulated to resist drought stress in purple wheat

LI Xiao-lan, Lü Xiang, WANG Xiao-hong, PENG Qin, ZHANG Ming-sheng, REN Ming-jian

2020, 19 (1): 33-50. DOI: 10.1016/S2095-3119(19)62659-6

Abstract （131） PDF in ScienceDirect

Triticum aestivum L. cv. Guizi 1 (GZ1) is a drought-tolerant local purple wheat cultivar. It is not clear how purple wheat resists drought stress, but it could be related to anthocyanin biosynthesis. In this study, transcriptome data from drought-treated samples and controls were compared. Drought slightly reduced the anthocyanin, protein and starch contents of GZ1 grains and significantly reduced the grain weight. Under drought stress, 16 682 transcripts were reduced, 27 766 differentially expressed genes (DEGs) were identified, and 379 DEGs, including DREBs, were related to defense response. The defense-response genes included response to water deprivation, reactive oxygen, bacteria, fungi, etc. Most of the structural and regulatory genes in anthocyanin biosynthesis were downregulated, with only TaDFR, TaOMT, Ta5,3GT, and TaMYB-4B1 being upregulated. TaCHS, TaF3H, TaCHI, Ta4CL, and TaF3’H are involved in responses to UV, hormones, and stimulus. TaCHS-2D1, TaDFR-2D2, TaDFR-7D, TaOMT-5A, Ta5,3GT-1B1, Ta5,3GT-3A, and Ta5,3GT-7B1 connect anthocyanin biosynthesis with other pathways, and their interacting proteins are involved in primary metabolism, genetic regulation, growth and development, and defense responses. There is further speculation about the defense-responsive network in purple wheat. The results indicated that biotic and abiotic stress-responsive genes were stimulated to resist drought stress in purple wheat GZ1, and anthocyanin biosynthesis also participated in the drought defense response through several structural genes.

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