Please wait a minute...
Journal of Integrative Agriculture  2022, Vol. 21 Issue (11): 3131-3147    DOI: 10.1016/j.jia.2022.08.085
Special Issue: 麦类遗传育种合辑Triticeae Crops Genetics · Breeding · Germplasm Resources
Crop Science Advanced Online Publication | Current Issue | Archive | Adv Search |
Identification and validation of novel loci associated with wheat quality through a genome-wide association study

PU Zhi-en1*, YE Xue-ling3, 4*, LI Yang1, SHI Bing-xin1, GUO Zhu1, DAI Shou-fen2, MA Jian2, LIU Ze-hou4, JIANG Yun-feng2, LI Wei1, JIANG Qian-tao2, CHEN Guo-yue2, WEI Yu-ming2, ZHENG You-liang2 #br#

1 College of Agronomy, Sichuan Agricultural University, Chengdu 611130, P.R.China

2 Triticeae Research Institute, Sichuan Agricultural University, Wenjiang 611130, P.R.China

3 Key Laboratory of Coarse Cereal Processing, Ministry of Agriculture and Rural Affairs/College of Food and Biological Engineering, Chengdu University, Chengdu 610106, P.R.China

4 Crop Research Institute, Sichuan Academy of Agricultural Sciences, Chengdu 610066, P.R.China

Download:  PDF in ScienceDirect  
Export:  BibTeX | EndNote (RIS)      
摘要  了解小麦品质相关性状的遗传基础有助于对小麦品质进行改良,本实验测定了多环境下236份小麦种质资源(包括 160 个栽培品种和76个地方品种)的蛋白质含量(GPC)、淀粉含量(GSC)和湿面筋含量(WGC),并使用 55K小麦芯片进行了混合线性模型 (MLM)分析。结果共鉴定了 12 个稳定的 QTL/SNP,与GPC、GSC和WGC 相关的位点分别有3个、7个和2个 QTL,它们分别位于1B、1D、2A、2B、2D、3B、3D、5D 和 7D 染色体上;表型变异解释 (PVE) 范围从4.2 至10.7%。与之前报道的 QTL/基因相比,5 个 QTL(QGsc.sicau-1BLQGsc.sicau-1DSQGsc.sicau-2DL.1QGsc.sicau-2DL.2QWgc.sicau-5DL)是潜在的新位点。本实验着重关注了位于5D染色体上与湿面筋浓度相关的稳定QTL,并成功开发了SNP AX-108770574AX-108791420 两个KASP 标记。其中AX-108770574中含有A-等位基因和AX-108791420中含有T-等位基因的品种表型显着高于(P<0.01)含有湿面筋浓度G-等位基因或C-等位基因的地方品种,表明开发的KASP 标记可用于分子育种,改良小麦品质。

Abstract  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.
Keywords:  cultivars        landraces        grain protein concentration (GPC)        grain starch concentration (GSC)        wet gluten concentration (WGC)        55K SNP       validation  
Received: 25 April 2021   Accepted: 01 July 2021
Fund: This work was supported by the National Key Research and Development Program of China (2017YFD0100900, 2016YFD0102000 and 2016YFD0100100), the International Science and Technology Cooperation and Exchanges Programs of Science and Technology Department of Sichuan Province, China (2019YFH0063), and the Sichuan Science and Technology Program, China (2022ZDZX0014).  
About author:  Correspondence ZHENG You-liang, Tel/Fax: +86-28-86290909, E-mail: ylzheng@sicau.edu.cn * These authors contributed equally to this study.

Cite this article: 

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. Identification and validation of novel loci associated with wheat quality through a genome-wide association study. Journal of Integrative Agriculture, 21(11): 3131-3147.

AACC (American Association of Cereal Chemists). 2000. Wet gluten, dry gluten, water-binding capacity and gluten index. International Method 38-12.02. USA.  
Appels R, Eversole K, Stein N, Feuillet C, Keller B, Rogers J, Pozniak CJ, Choulet F, Distelfeld A, Poland J, Ronen G. 2018. Shifting the limits in wheat research and breeding using a fully annotated reference genome. Science, 361, 7191.
Avci U, Earl Petzold H, Ismail I O, Beers E P, Haigler C H. 2008. Cysteine proteases XCP1 and XCP2 aid micro-autolysis within the intact central vacuole during xylogenesis in Arabidopsis roots. The Plant Journal, 56, 303–315.
Balyan H S, Gupta P K, Kumar S, Dhariwal R, Jaiswal V, Tyagi S, Agarwal P, Gahlaut V, Kumari S. 2013. Genetic improvement of grain protein content and other health-related constituents of wheat grain. Plant Breeding, 132, 446–457.
Bates D, Maechler M, Bolker B, Walker S. 2014. LME4: Linear mixed - Effects models using eigen and S4. R Package version 1.1-4.
Bazakos C, Hanemian M, Trontin C, Jiménez-Gómez J M, Loudet O. 2017. New strategies and tools in quantitative genetics: How to go from the phenotype to the genotype. Annual Review of Plant Biology, 68, 435–455.
Bogard M, Allard V, Martre P, Heumez E, Snape J W, Orford S, Griffiths S, Gaju O, Foulkes J, Le Gouis J. 2013. Identifying wheat genomic regions for improving grain protein concentration independently of grain yield using multiple inter-related populations. Molecular Breeding, 31, 587–599.
Bradbury P J, Zhang Z, Kroon D E, Casstevens T M, Ramdoss Y, Buckler E S. 2007. TASSEL: Software for association mapping of complex traits in diverse samples. Bioinformatics, 23, 2633–2635.
Chakraborty R, Jin L. 1993. A unified approach to study hypervariable polymorphisms: Statistical considerations of determining relatedness and population distances. In: DNA Fingerprinting: State of the Science. Birkhäuser, Basel. pp. 153–175. 
Chen J, Zhang F, Zhao C, Lv G, Sun C, Pan Y, Guo X, Chen F. 2019. Genome-wide association study of six quality traits reveals the association of the TaRPP13L1 gene with flour colour in Chinese bread wheat. Plant Biotechnology Journal, 17, 2106–2122.
Chia T, Chirico M, King R, Ramirez-Gonzalez R, Saccomanno B, Seung D, Simmonds J, Trick M, Uauy C, Verhoeven T, Trafford K. 2020. A carbohydrate-binding protein, B-GRANULE CONTENT 1, influences starch granule size distribution in a dose-dependent manner in polyploid wheat. Journal of Experimental Botany, 71, 105–115.
Chiotelli E, Le Meste M. 2002. Effect of small and large wheat starch granules on thermomechanical behavior of starch. Cereal Chemistry, 79, 286–293.
Cui F, Fan X, Chen M, Zhang N, Zhao C, Zhang W, Han J, Ji J, Zhao X, Yang L, Zhao Z. 2016. QTL detection for wheat kernel size and quality and the responses of these traits to low nitrogen stress. Theoretical and Applied Genetics, 129, 469–484.
DalCorso G, Pesaresi P, Masiero S, Aseeva E, Schünemann D, Finazzi G, Joliot P, Barbato R, Leister D. 2008. A complex containing PGRL1 and PGR5 is involved in the switch between linear and cyclic electron flow in Arabidopsis. Cell, 132, 273–285.
Denčić S R, Mladenov N O, Kobiljski B O. 2012. Effects of genotype and environment on bread making quality in wheat. International Journal of Plant Production, 5, 71–82.
Deng Z, Fang W, Guo X, Zhao X, Guo H, Hu S, Tian J. 2018. Genetic dissection of interactions between wheat flour starch and its components in two populations using two QTL mapping methods. Molecular Breeding, 38, 41.
Deng Z, Hu S, Chen F, Li W, Chen J, Sun C, Zhang Y, Wang S, Song X, Tian J. 2015a. Genetic dissection of interaction between wheat protein and starch using three mapping populations. Molecular Breeding, 35, 12.
Deng Z, Tian J, Chen F, Li W, Zheng F, Chen J, Shi C, Sun C, Wang S, Zhang Y. 2015b. Genetic dissection on wheat flour quality traits in two related populations. Euphytica, 203, 221–235.
Deng Z Y, Zhao L, Liu B, Zhang K P, Chen J S, Qu H L, Sun C L, Zhang Y X, Tian J C. 2013. Conditional QTL mapping of sedimentation volume on seven quality traits in common wheat. Journal of Integrative Agriculture, 12, 2125–2133.
Devadas R, Simpfendorfer S, Backhouse D, Lamb D W. 2014. Effect of stripe rust on the yield response of wheat to nitrogen. The Crop Journal, 2, 201–206.
Dinant S, Clark A M, Zhu Y, Vilaine F, Palauqui J C, Kusiak C, Thompson G A. 2003. Diversity of the superfamily of phloem lectins (phloem protein 2) in angiosperms. Plant Physiology, 131, 114–128.
Earl D A. 2012. STRUCTURE HARVESTER: A website and program for visualizing STRUCTURE output and implementing the Evanno method. Conservation Genetics Resources, 4, 359–361.
Echeverry-Solarte M, Kumar A, Kianian S, Simsek S, Alamri M S, Mantovani E E, McClean P E, Deckard E L, Elias E, Schatz B, Xu S S. 2015. New QTL alleles for quality-related traits in spring wheat revealed by RIL population derived from supernumerary×non-supernumerary spikelet genotypes. Theoretical and Applied Genetics, 128, 893–912.
Fan X, Cui F, Ji J, Zhang W, Zhao X, Liu J, Tong Y, Wang T, Li J. 2019. Dissection of pleiotropic QTL regions controlling wheat spike characteristics under different nitrogen treatments using traditional and conditional QTL mapping. Frontier in Plant Science, 10, 187. 
Fatiukha A, Filler N, Lupo I, Lidzbarsky G, Klymiuk, V, Korol A B, Pozniak C, Fahima T, Krugman T. 2020. Grain protein content and thousand kernel weight QTLs identified in a durum×wild emmer wheat mapping population tested in five environments. Theoretical and Applied Genetics, 133, 119–131.
Fernando N, Panozzo J, Tausz M, Norton R, Fitzgerald G, Seneweera S. 2012. Rising atmospheric CO2 concentration affects mineral nutrient and protein concentration of wheat grain. Food Chemistry, 133, 1307–1311.
Fiedler J D, Salsman E, Liu Y, Michalak de Jiménez M, Hegstad J B, Chen B, Manthey F A, Chao S, Xu S, Elias E M, Li X. 2017. Genome-wide association and prediction of grain and semolina quality traits in durum wheat breeding populations. The Plant Genome, 10, 1–12.
García-Cerdán J G, Kovács L, Tóth T, Kereïche S, Aseeva E, Boekema E J, Mamedov F, Funk C, Schröder W P. 2011. The PsbW protein stabilizes the supramolecular organization of photosystem II in higher plants. The Plant Journal, 65, 368–381.
Gaudet P, Livstone M S, Lewis S E, Thomas P D. 2011. Phylogenetic-based propagation of functional annotations within the Gene Ontology consortium. Briefings in Bioinformatics, 12, 449–462.
Gillies S A, Futardo A, Henry R J. 2012. Gene expression in the developing aleurone and starchy endosperm of wheat. Plant Biotechnology Journal, 10, 668–679.
Goel S, Singh K, Singh B, Grewal S, Dwivedi N, Alqarawi A A, Abd Allah E F, Ahmad P, Singh N K. 2019. Analysis of genetic control and QTL mapping of essential wheat grain quality traits in a recombinant inbred population. PLoS ONE, 14, e0200669.
Grimaud F, Rogniaux H, James M G, Myers A M, Planchot V. 2008. Proteome and phosphoproteome analysis of starch granule-associated proteins from normal maize and mutants affected in starch biosynthesis. Journal of Experimental Botany, 59, 3395–3406.
Groos C, Robert N, Bervas E, Charmet G. 2003. Genetic analysis of grain protein-content, grain yield and thousand-kernel weight in bread wheat. Theoretical and Applied Genetics, 106, 1032–1040.
Guan X. 2018. Genome-wide association analysis of main starch quality and dough rheological properties in wheat. MSc thesis, Shandong Agricultural University, China. (in Chinese)
Haynes L C, Bettge A D, Slade L. 2009. Soft wheat and flour products methods review: Solvent retention capacity equation correction. Cereal Foods World, 54, 174–175.
Hertle A P, Blunder T, Wunder T, Pesaresi P, Pribil M, Armbruster U, Leister D. 2013. PGRL1 is the elusive ferredoxin-plastoquinone reductase in photosynthetic cyclic electron flow. Molecular Cell, 49, 511–523.
Hu S N. 2013. Conditional QTL analysis of interaction between wheat protein and starch content. MSc thesis, Shandong Agricultural University, China. (in Chinese)
Jernigan K L, Godoy J V, Huang M, Zhou Y, Morris C F, Garland-Campbell K A, Zhang Z, Carter A H. 2018. Genetic dissection of end-use quality traits in adapted soft white winter wheat. Frontiers in Plant Science, 9, 271.
Johnson M, Kumar A, Oladzad-Abbasabadi A, Salsman E, Aoun M, Manthey F A, Elias E M. 2019. Association mapping for 24 traits related to protein content, gluten strength, color, cooking and milling quality using balanced and unbalanced data in durum wheat [Triticum Turgidum L. var. Durum (Desf.)]. Frontiers in Genetics, 10, 717.
Juliana P, Poland J, Huerta-Espino J, Shrestha S, Crossa J, Crespo-Herrera L, Toledo F H, Govindan V, Mondal S, Kumar U, Bhavani S. 2019. Improving grain yield, stress resilience and quality of bread wheat using large-scale genomics. Nature Genetics, 51, 1530–1539.
Kim T H, Kim B H, Yahalom A, Chamovitz D A, von Arnim A G. 2004. Translational regulation via 5´ mRNA leader sequences revealed by mutational analysis of the Arabidopsis translation initiation factor subunit eIF3h. The Plant Cell, 16, 3341–3356.
Kodama A, Narita R, Yamaguchi M, Hisano H, Adachi S, Takagi H, Ookawa T, Sato K, Hirasawa T. 2018. QTLs maintaining grain fertility under salt stress detected by exome QTL-seq and interval mapping in barley. Breeding Science, 68, 561–570.
Kong L G, Si J S, Zhang B, Feng B, Li S D, Wang F H. 2013. Environmental modification of wheat grain protein accumulation and associated processing quality: a case study of china. Australian Journal of Crop Science, 7, 173–181.
Krystkowiak K, Langner M, Adamski T, Salmanowicz B P, Kaczmarek Z, Krajewski P, Surma M. 2017. Interactions between Glu-1 and Glu-3 loci and associations of selected molecular markers with quality traits in winter wheat (Triticum aestivum L.) DH lines. Journal of Applied Genetics, 58, 37–48.
Kumar A, Jain S, Elias E M, Ibrahim M, Sharma L K. 2018. An overview of QTL identification and marker-assisted selection for grain protein content in wheat. In: Eco-Friendly Agro-Biological Techniques for Enhancing Crop Productivity. Springer, Singapore. pp. 245–274. 
Labuschagne M T, Geleta N, Osthoff G. 2007. The influence of environment on starch content and amylose to amylopectin ratio in wheat. Starch-Stärke, 59, 234–238.
Letunic I, Bork P. 2019. Interactive Tree of life (iTOL) v4: Recent updates and new developments. Nucleic Acids Research, 47, W256–W259.
Li C, Huang Y, Huang R, Wu Y, Wang W. 2018. The genetic architecture of amylose biosynthesis in maize kernel. Plant Biotechnology Journal, 16, 688–695.
Li H M, Tang Z X, Zhang H Q, Yan B J, Ren Z L. 2013. Major quality trait analysis and QTL detection in hexaploid wheat in humid rain-fed agriculture. Genetics and Molecular Research, 12, 1740–1751.
Li J, Cui F, Ding A M, Zhao C H, Wang X Q, Wang L, Bao Y G, Qi X L, Li X F, Gao J R, Feng D S. 2012. QTL detection of seven quality traits in wheat using two related recombinant inbred line populations. Euphytica, 183, 207–226.
Li R, Zeng Y, Xu J, Wang Q, Wu F, Cao M, Lan H, Liu Y, Lu Y. 2015. Genetic variation for maize root architecture in response to drought stress at the seedling stage. Breeding Science, 65, 298–307.
Li Y, Song Y, Zhou R, Branlard G, Jia J. 2009. Detection of QTLs for bread-making quality in wheat using a recombinant inbred line population. Plant Breeding, 128, 235–243.
Liu J, Feng B, Xu Z, Fan X, Jiang F, Jin X, Cao J, Wang F, Liu Q, Yang L, Wang T. 2018. A genome-wide association study of wheat yield and quality-related traits in southwest China. Molecular Breeding, 38, 1.
Liu J, Wu B, Singh R P, Velu G. 2019. QTL mapping for micronutrients concentration and yield component traits in a hexaploid wheat mapping population. Journal of Cereal Science, 88, 57–64.
Liu K, Muse S V. 2005. PowerMarker: an integrated analysis environment for genetic marker analysis. Bioinformatics, 21, 2128–2129.
Liu T, An Y, Liu K, Wang F, Xie C, Zhang Y, Guan X, Tian J, Chen J. 2017. A genetic analysis of the quality of northern-style Chinese steamed bread. Molecular Breeding, 37, 41.
Liu T T, Xu M Z, Gao S Q, Zhang Y, Hu Y, Jin P, Cai L N, Cheng Y, Chen J P, Yang J, Zhong K L. 2022. Genome-wide identification and analysis of the regulation wheat DnaJ family genes following wheat yellow mosaic virus infection. Journal of Integrative Agriculture, 21, 153–169.
Ługowska B, Obuchowskił W, Salmanowicz B, Krystkowiak K. 2012. Effect of genotype, environment and their interaction on quality parameters of wheat breeding lines of diverse grain hardness. Plant Production Science, 15, 192–203.
Ma Y M, Lou H Y, Chen Z Y, Xiao J, Xu L, Ni Z F, Liu J. 2020. Genetic diversity assessment of winter wheat landraces and cultivars in Xinjiang via SNP array analysis. Acta Agronomica Sinica, 46, 1539–1556. (in Chinese)
Marcotuli I, Gadaleta A, Mangini G, Signorile A M, Zacheo S A, Blanco A, Simeone R, Colasuonno P. 2017. Development of a high-density SNP-based linkage map and detection of QTL for β-glucans, protein content, grain yield per spike and heading time in durum wheat. International Journal of Molecular Sciences, 18, 1329.
McCartney C A, Somers D J, Lukow O, Ames N, Noll J, Cloutier S, Humphreys D G, McCallum B D. 2006. QTL analysis of quality traits in the spring wheat cross RL4452בAC Domain’. Plant Breeding, 125, 565–575.
Mir Drikvand R, Najafian G, Bihamta M R, Ebrahimi A. 2018. Mapping some seed quality traits in bread wheat (Triticum aestivum L.) by association mapping using SSR markers. Journal of Applied Biotechnology Reports, 5, 92–99.
Muhammad S, Sajjad M, Khan S H, Shahid M, Zubair M, Awan F S, Khan A I, Mubarak M S, Tahir A, Umer M, Keyani R, Afzal M I, Manzoor I, Wattoo J I, Rehman A. 2020. Genome-wide association analysis for stripe rust resistance in spring wheat (Triticum aestivum L.) germplasm. Journal of Integrative Agriculture, 19, 2035–2043.
Nedelkou I P, Maurer A, Schubert A, Léon J, Pillen K. 2017. Exotic QTL improve grain quality in the tri-parental wheat population SW84. PLoS ONE, 12, e0179851.
Nigro D, Gadaleta A, Mangini G, Colasuonno P, Marcotuli I, Giancaspro A, Giove S L, Simeone R, Blanco A. 2019. Candidate genes and genome-wide association study of grain protein content and protein deviation in durum wheat. Planta, 249, 1157–1175.
O’Brien L, Brown J S, Panozzo J F, Archer M J. 1990. The effect of stripe rust on the quality of Australian wheat varieties. Australian Journal of Agricultural Research, 41, 827–833.
Pang H, Wang L, Wang H L, Xu H J, Li W H. 2014. QTL mapping for kernel starch content in wheat. Journal of Triticeae Crops, 34, 1–7. (in Chinese)
Panozzo J F, McCormick K M. 1993. The rapid viscoanalyser as a method of testing for noodle quality in a wheat breeding programme. Journal of Cereal Science, 17, 25–32.
Preiss J. 1991. Biology and molecular biology of starch synthesis and its regulation. In: BJ Miflin B J, ed., Oxford Surveys of Plant Molecular and Cell Biology. vol. 7. Oxford University Press, Oxford, UK. pp. 59–114.
Pritchard J K, Stephens M, Donnelly P. 2000. Inference of population structure using multilocus genotype data. Genetics, 155, 945–959.
Qi H, Xia F N, Xie L J, Yu L J, Chen Q F, Zhuang X H, Wang Q, Li F, Jiang L, Xie Q, Xiao S. 2017. TRAF family proteins regulate autophagy dynamics by modulating AUTOPHAGY PROTEIN6 stability in Arabidopsis. The Plant Cell, 29, 890–911.
Rakszegi M, Mikó P, Löschenberger F, Hiltbrunner J, Aebi R, Knapp S, Tremmel-Bede K, Megyeri M, Kovács G, Molnár-Láng M, Vida G. 2016. Comparison of quality parameters of wheat varieties with different breeding origin under organic and low-input conventional conditions. Journal of Cereal Science, 69, 297–305.
Rapp M, Lein V, Lacoudre F, Lafferty J, Müller E, Vida G, Bozhanova V, Ibraliu A, Thorwarth P, Piepho H P, Leiser W L. 2018. Simultaneous improvement of grain yield and protein content in durum wheat by different phenotypic indices and genomic selection. Theoretical and Applied Genetics, 131, 1315–1329.
Reif J C, Gowda M, Maurer H P, Longin C F, Korzun V, Ebmeyer E, Bothe R, Pietsch C, Würschum T. 2011. Association mapping for quality traits in soft winter wheat. Theoretical and Applied Genetics, 122, 961–970.
Roselló M, Royo C, Álvaro F, Villegas D, Nazco R, Soriano J M. 2018. Pasta-making quality QTLome from Mediterranean durum wheat landraces. Frontiers in Plant Science, 9, 1512.
Sado P E, Tessier D, Vasseur M, Elmorjani K, Guillon F, Saulnier L. 2009. Integrating genes and phenotype: A wheat–Arabidopsis–rice glycosyltransferase database for candidate gene analyses. Functional & Integrative Genomics, 9, 43–58.
Simmonds N W. 1995. The relation between yield and protein in cereal grain. Journal of the Science of Food and Agriculture, 67, 309–315.
Smith S E, Kuehl R O, Ray I M, Hui R, Soleri D. 1998. Evaluation of simple methods for estimating broad-sense heritability in stands of randomly planted genotypes. Crop Science, 38, 1125–1129.
Su Y, Liao P, Song D, Huang S, He J, Gao X, Li S. 2020. Application of Aegilops tauschii–Triticum aestivum recombinant inbred lines for grain protein content QTL detection and wheat improvement. Canadian Journal of Plant Science, 100, 425–434.
Sukumaran S, Reynolds M P, Sansaloni C. 2018. Genome-wide association analyses identify QTL hotspots for yield and component traits in durum wheat grown under yield potential, drought, and heat stress environments. Frontiers in Plant Science, 9, 81. 
Sunderman D W, Wise M. 1964. Influence of stripe rust of wheat upon plant development and grain quality of closely related Lemhi derivatives 1. Crop Science, 4, 347–348.
Surma M, Adamski T, Banaszak Z, Kaczmarek Z, Kuczyńska H, Majcher M, Ługowska B, Obuchowskił W, Salmanowicz B, Krystkowiak K. 2012. Effect of genotype, environment and their interaction on quality parameters of wheat breeding lines of diverse grain hardness. Plant Production Science, 15, 192–203.
Tabbita F, Pearce S, Barneix A J. 2017. Breeding for increased grain protein and micronutrient content in wheat: Ten years of the GPC-B1 gene. Journal of Cereal Science, 73, 183–191.
Thorwarth P, Liu G, Ebmeyer E, Schacht J, Schachschneider R, Kazman E, Reif J C, Würschum T, Longin C F. 2019. Dissecting the genetics underlying the relationship between protein content and grain yield in a large hybrid wheat population. Theoretical and Applied Genetics, 132, 489–500.
Tian B, Deng Z, Xie Q, Tian J. 2015. Genetic dissection of the developmental behaviour of total starch content and its components in wheat grain. Crop and Pasture Science, 66, 445–455.
Tian J, Chen J, Chen G, Wu P, Zhang H, Zhao Y. 2015a. Conditional QTL mapping of major quality traits. In: Genetic Analyses of Wheat and Molecular Marker-Assisted Breeding. vol. 2. Springer, Dordrecht. pp. 15–88.
Tian J, Deng Z, Zhang K, Yu H, Jiang X, Li C. 2015b. Genetic Detection of main quality traits in wheat. In: Genetic Analyses of Wheat and Molecular Marker-Assisted Breeding. vol. 1. Springer, Dordrecht. pp. 177–350. 
Torii K U, Mitsukawa N, Oosumi T, Matsuura Y, Yokoyama R, Whittier R F, Komeda Y. 1996. The Arabidopsis ERECTA gene encodes a putative receptor protein kinase with extracellular leucine-rich repeats. The Plant Cell, 8, 735–746.
Turner A S, Bradburne R P, Fish L, Snape J W. 2004. New quantitative trait loci influencing grain texture and protein content in bread wheat. Journal of Cereal Science, 40, 51-60.
Wickham H. 2016. Ggplot2: Elegant Graphics for Data Analysis. Springer, American. 
Willmann M R, Mehalick A J, Packer R L, Jenik P D. 2011. MicroRNAs regulate the timing of embryo maturation in Arabidopsis. Plant Physiology, 155, 1871–1884.
Ye X, Li J, Cheng Y, Yao F, Long L, Wang Y, Wu Y, Li J, Wang J, Jiang Q, Kang H. 2019a. Genome-wide association study reveals new loci for yield-related traits in Sichuan wheat germplasm under stripe rust stress. BMC Genomics, 20, 1–17.
Ye X, Li J, Cheng Y, Yao F, Long L, Yu C, Wang Y, Wu Y, Li J, Wang J, Jiang Q. 2019b. Genome-wide association study of resistance to stripe rust (Puccinia striiformis f. sp. tritici) in Sichuan wheat. BMC Plant Biology, 19, 1–5.
Young T E, Gallie D R. 1999. Analysis of programmed cell death in wheat endosperm reveals differences in endosperm development between cereals. Plant Molecular Biology, 39, 915–926.
Zhang C C. 2019. Genome-wide association analysis of flour processing quality traits related of Triticum dicoccoides and cloning of high molecular weight glutenin subunit gene 1AX1. MSc thesis, Henan University, China. (in Chinese)
Zhang W, Chao S, Manthey F, Chicaiza O, Brevis J C, Echenique V, Dubcovsky J. 2008. QTL analysis of pasta quality using a composite microsatellite and SNP map of durum wheat. Theoretical and Applied Genetics, 117, 1361–1377.
Zhang Y F. 2016. Research on the effects of high temperature stress on starch and protein content of wheat grains. Ph D thesis, China Agricultural University, China. (in Chinese)
Zhou Y, Li S, Qian Q, Zeng D, Zhang M, Guo L, Liu X, Zhang B, Deng L, Liu X, Luo G. 2009. BC10, a DUF266-containing and Golgi-located type II membrane protein, is required for cell-wall biosynthesis in rice (Oryza sativa L.). The Plant Journal, 57, 446–462.
Zou J, Semagn K, Iqbal M, N’Diaye A, Chen H, Asif M, Navabi A, Perez-Lara E, Pozniak C, Yang R C, Randhawa H. 2017. Mapping QTLs controlling agronomic traits in the ‘Attila’בCDC Go’ spring wheat population under organic management using 90K SNP array. Crop Science, 57, 365–377.


[1] WU Xian-xin, ZANG Chao-qun, ZHANG Ya-zhao, XU Yi-wei, WANG Shu, LI Tian-ya, GAO Li.

Characterization of wheat monogenic lines with known Sr genes and wheat cultivars for resistance to three new races of Puccinia graminis f. sp. tritici in China [J]. >Journal of Integrative Agriculture, 2023, 22(6): 1740-1749.

[2] MA Da-ling, XIE Rui-zhi, YU Xiao-fang, LI Shao-kun, GAO Ju-lin. Historical trends in maize morphology from the 1950s to the 2010s in China[J]. >Journal of Integrative Agriculture, 2022, 21(8): 2159-2167.
[3] PAN Wen-jing, HAN Xue, HUANG Shi-yu, YU Jing-yao, ZHAO Ying, QU Ke-xin, ZHANG Ze-xin, YIN Zhen-gong, QI Hui-dong, YU Guo-long, ZHANG Yong, XIN Da-wei, ZHU Rong-sheng, LIU Chun-yan, WU Xiao-xia, JIANG Hong-wei, HU Zhen-bang, ZUO Yu-hu, CHEN Qing-shan, QI Zhao-ming. Identification of candidate genes related to soluble sugar contents in soybean seeds using multiple genetic analyses[J]. >Journal of Integrative Agriculture, 2022, 21(7): 1886-1902.
[4] 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. A major and stable QTL for wheat spikelet number per spike validated in different genetic backgrounds[J]. >Journal of Integrative Agriculture, 2022, 21(6): 1551-1562.
[5] ZHANG Li-xin, LIU Wei, Mesfin Tsegaw, XU Xin, QI Yan-ping, Enoch Sapey, LIU Lu-ping, WU Ting-ting, SUN Shi, HAN Tian-fu. Principles and practices of the photo-thermal adaptability improvement in soybean[J]. >Journal of Integrative Agriculture, 2020, 19(2): 295-310.
[6] HUANG Cheng-dong, LIU Quan-qing, LI Xiao-lin, ZHANG Chao-chun. Effect of intercropping on maize grain yield and yield components[J]. >Journal of Integrative Agriculture, 2019, 18(8): 1690-1700.
[7] ZHOU Liang, MU Tai-hua, MA Meng-mei, ZHANG Ruo-fang, SUN Qing-hua, XU Yan-wen. Nutritional evaluation of different cultivars of potatoes (Solanum tuberosum L.) from China by grey relational analysis (GRA) and its application in potato steamed bread making[J]. >Journal of Integrative Agriculture, 2019, 18(1): 231-245.
[8] LI Hui-xia, CHEN Zhu-jun, ZHOU Ting, LIU Yan, ZHOU Jian-bin. High potassium to magnesium ratio affected the growth and magnesium uptake of three tomato (Solanum lycopersicum L.) cultivars[J]. >Journal of Integrative Agriculture, 2018, 17(12): 2813-2821.
[9] CHANG Li-fang, LI Hui-hui, WU Xiao-yang, LU Yu-qing, ZHANG Jin-peng, YANG Xin-ming, LI Xiu-quan, LIU Wei-hua, LI Li-hui. Genetic characteristics of a wheat founder parent and a widely planted cultivar derived from the same cross[J]. >Journal of Integrative Agriculture, 2018, 17(04): 775-785.
[10] HE Ying-bin, ZHOU Yang-fan, CAI Wei-min, WANG Zhuo-zhuo, DUAN Ding-ding, LUO Shan-jun, CHEN Jing-zhu. Using a process-oriented methodology to precisely evaluate temperature suitability for potato growth in China using GIS[J]. >Journal of Integrative Agriculture, 2017, 16(07): 1520-1529.
[11] ZHU Bo, ZHANG Jing-jing, NIU Hong, GUAN Long, GUO Peng, XU Ling-yang, CHEN Yan, ZHANG Lu-pei, GAO Hui-jiang, GAO Xue, LI Jun-ya. Effects of marker density and minor allele frequency on genomic prediction for growth traits in Chinese Simmental beef cattle[J]. >Journal of Integrative Agriculture, 2017, 16(04): 911-920.
[12] AO Yan, XU Yong, CUI Xiao-fen, WANG An, TENG Fei, SHEN Li-qun, LIU Qiao-quan. A genetic diversity assessment of starch quality traits in rice landraces from the Taihu basin, China[J]. >Journal of Integrative Agriculture, 2016, 15(3): 493-501.
[13] WANG Yun-qi, XI Wen-xing, WANG Zhi-min, WANG Bin, XU Xue-xin, HAN Mei-kun, ZHOU Shun-li, ZHANG Ying-hua. Contribution of ear photosynthesis to grain yield under rainfed and irrigation conditions for winter wheat cultivars released in the past 30 years in North China Plain[J]. >Journal of Integrative Agriculture, 2016, 15(10): 2247-2256.
[14] Jan Bocianowski, Piotr Szulc, Anna Tratwal, Kamila Nowosad, Dariusz Piesik. The influence of potassium to mineral fertilizers on the maize health[J]. >Journal of Integrative Agriculture, 2016, 15(06): 1286-1292.
[15] SONG Wen-en, CHEN Shi-bao, LIU Ji-fang, CHEN Li, SONG Ning-ning, LI Ning, LIU Bin. Variation of Cd concentration in various rice cultivars and derivation of cadmium toxicity thresholds for paddy soil by species-sensitivity distribution[J]. >Journal of Integrative Agriculture, 2015, 14(9): 1845-1854.
No Suggested Reading articles found!