Allen G C, Flores-Vergara M A, Krasynanski S, Kumar S, Thompson W F. 2006. A modified protocol for rapid DNA isolation from plant tissues using cetyltrimethylammonium bromide. Nature Protocols, 1, 2320–2325.

Anjum F M, Ahmad I, Butt M S, Sheikh M A, Pasha I. 2005. Amino acid composition of spring wheats and losses of lysine during chapati baking. Journal of Food Composition & Analysis, 18, 523–532.

Arjinajarn P, Chueakula N, Pongchaidecha A, Jaikumkao K, Chatsudthipong V, Mahatheeranont S, Norkaew O, Chattipakorn N, Lungkaphin A. 2017. Anthocyanin-rich riceberry bran extract attenuates gentamicin-induced hepatotoxicity by reducing oxidative stress, inflammation and apoptosis in rats. Biomedicine Pharmacotherapy, 92, 412–420.

Bai L. 2012. Comparative mapping of wheat–Agropyron cristatum 6P addition lines and molecular markers of its introgression lines. MSc thesis, Chinese Academy of Agricultural Sciences, Beijing. (in Chinese)

Bolger A M, Lohse M, Usadel B. 2014. Trimmomatic: A flexible trimmer for Illumina sequence data. Bioinformatics, 30, 2114–2120.

Bondonno N P, Dalgaard F, Kyrø C, Murray K, Bondonno C P, Lewis J R, Croft K D, Gislason G, Scalbert A, Cassidy A, Tjønneland A, Overvad K, Hodgson J M. 2019. Flavonoid intake is associated with lower mortality in the Danish diet cancer and health cohort. Nature Communications, 10, 3651.

Cao L, Han G H, Liu H, Yan H W, Jin Y L, Cao L J, Zhou Y L, An D G. 2023. Development of novel wheat–rye 6RS small fragment translocation lines with powdery mildew resistance and physical mapping of the resistance gene PmW6RS. Theoretical and Applied Genetics, 136, 108.

Chen D, Zhang J P, Wang J S, Yang X M, Liu W H, Gao A N, Li X Q, Li L H. 2012. Inheritance and availability of high grain number per spike in two wheat germplasm lines. Journal of Integrative Agriculture, 11, 1409–1416.

Chen H Y, Chen X D, Yu L S. 2020. Research progress on the formation mechanism, biological function and application development of melanin. Biological Resources, 42, 652–659.

Cheng X F, Xiao Y, Wang L H, Yang X Y, Deng P C, Zhao J X, Wang C Y, Chen C H, Li T D, Ji W Q. 2026. Cytogenetic characterization and molecular marker development of a novel wheat–Thinopyrum ponticum 5E (5D) disomic substitution line with resistance to powdery mildew and stripe rust. Journal of Integrative Agriculture, 25, 30–41.

Dong Y C, Zheng D S. 2000. Genetic Resources of Wheat in China. China Agricultural Press, Beijing. p. 175.

Du H M, Tang Z X, Duan Q, Tang S Y, Fu S L. 2018. Using the 6RL^ku minichromosome of rye (Secale cereale L.) to create wheat–rye 6D/6RL^ku small segment translocation lines with powdery mildew resistance. International Journal of Molecular Sciences, 19, 3933.

Friebe B, Kynast R G, Zhang P, Qi L L, Dhar M, Gill B S. 2001. Chromosome healing by addition of telomeric repeats in wheat occurs during the first mitotic divisions of the sporophyte and is a gradual process. Chromosome Research, 9, 137–146.

Friebe B, Larter E N. 1988. Identification of a complete set of isogenic wheat–rye D genome substitution lines by means of Giemsa C-banding. Theoretical and Applied Genetics, 76, 473–479.

Friebe B, Zhang P, Linc G, Gill B S. 2005. Robertsonian translocations in wheat arise by centric misdivision of univalents at anaphase I and rejoining of broken centromeres during interkinesis of meiosis II. Cytogenetic and Genome Research, 109, 293–297.

GB 5009.268-2016. 2016. National Food Safety Standard: Determination of Multi-elements in Food. National Health and Family Planning Commission of the People's Republic of China. Standards Press of China, Beijing.

Gordeeva E I, Glagoleva A Y, Kukoeva T V, Khlestkina E K, Shoevaet O Y. 2019. Purple-seeded barley (Hordeum vulgare L.): Marker-assisted development of NILs for investigating peculiarities of the anthocyanin biosynthesis regulatory network. BMC Plant Biology, 19, 49–57.

Gordeeva E I, Shoeva O Y, Khlestkina E K. 2015. Marker-assisted development of bread-wheat near-isogenic lines carrying various combinations of purple pericarp (Pp) alleles. Euphytica, 203, 469–476.

Guo Z F, Zhang Z B, Xu P, Guo Y N. 2013. Analysis of nutrient composition of purple wheat. Cereal Research Communications, 41, 293–303.

Han G H, Wang J, Yan H W, Cao L J, Liu S Y, Li X Q, Zhou Y L, Liu W, Gu T T, Shi Z P, Liu H, Li L H, An D G. 2025. Development and molecular cytogenetic identification of a new wheat–rye 6RL ditelosomic addition and 1R (1B) substitution line with powdery mildew resistance. Journal of Integrative Agriculture, 24, 72–84.

Han H M, Liu W H, Zhang J P, Zhou S H, Yang X M, Li X Q, Li L H. 2019a. Identification of P genome chromosomes in Agropyron cristatum and wheat–A.cristatum derivative lines by FISH. Scientific Reports, 9, 9712.

Han H M, Ma X Y, Wang Z, Qi K, Yang W J, Liu W H, Zhang J P, Zhou S H, Lu Y Q, Yang X M, Li X Q, Li L H. 2023. Chromosome 5P of Agropyron cristatum induces chromosomal translocation by disturbing homologous chromosome pairing in a common wheat background. Crop Journal, 11, 228–237.

Han H M, Zhang Y X, Liu W H, Hu Z M, Li L H. 2015. Degenerate oligonucleotide primed-polymerase chain reaction-based chromosome painting of P genome chromosomes in Agropyron cristatum and Wheat–A. cristatum addition lines. Crop Science, 55, 2798–2805.

Han R, Diao R N, Liang H F, Kuo Y, Wang X L. 2019b. Agropyron cristatum, an important gene source in wheat genetic improvement. Shandong Agricultural Sciences, 51, 1–9. (in Chinese)

He Z H, Zhuang Q S, Cheng S H, Yu Z W, Zhao Z D, Liu X. 2018. China’s wheat industry development and scientific and technological progress. Journal of Agricultural Sciences, 8, 99–106.

Huang W. 2012. Genetic diversity and genetic analysis of grain pigment content in colored wheat. MSc thesis, Shandong Agricultural University, China. (in Chinese)

Jiang B, Liu T G, Li H H, Han H M, Li L H, Zhang J P, Yang X M, Zhou S H, Li X Q, Liu W H. 2018a. Physical mapping of a novel locus conferring leaf rust resistance on the long arm of Agropyron cristatum chromosome 2P. Frontiers in Plant Science, 9, 817.

Jiang W, Liu T, Nan W, Jeewani D C, Niu Y, Li C, Wang Y, Shi X, Wang C, Wang J, Li Y, Gao X, Wang Z. 2018b. Two transcription factors TaPpm1 and TaPpb1 co-regulate the anthocyanin biosynthesis in purple pericarp of wheat. Journal of Experimental Botany, 69, 2555–2567.

Kang H Y, Zhang Z J, Xu L L, Qi W L, Tang Y, Wang H, Zhu W, Li D Y, Zeng J, Wang Y, Fan X, Sha L N, Zhang H Q, Zhou Y H. 2016. Characterization of wheat–Psathyrostachys huashanica small segment translocation line with enhanced kernels per spike and stripe rust resistance. Genome, 59, 221–229.

Khlestkina E K. 2013. Genes determining the coloration of different organs in wheat. Russian Journal of Genetics, 3, 54–65.

Kil Y S, Han A R, Hong M J, Kim J B, Park P H, Choi H, Nam J W. 2021. ¹H NMR-based chemometrics to gain insights into the bran of radiation-induced colored wheat mutant. Frontiers in Nutrition, 8, 806744.

Lachman J, Hejtmánková A, Orsák M, Popov M, Martinek P. 2017a. Tocotrienols and tocopherols in colored-grain wheat, tritordeum and barley. Food Chemistry, 240, 725–35.

Lachman J, Martinek P, Kotíková Z, Orsak M, Sulc M. 2017b. Genetics and chemistry of pigments in wheat seeds-A review. Journal of Cereal Science, 74, 145–154.

Lan S Q, Li X P, Liu Y P. 2008. Grain pigment inheritance of blue grain wheat. Journal of Agriculture of North China, 23, 12–14.

Levy A A, Feldman M. 2022. Evolution and origin of bread wheat. The Plant Cell, 34, 2549–2567.

Li H, Durbin R. 2009. Fast and accurate short read alignment with Burrows–Wheeler transform. Bioinformatics, 25, 1754–1760.

Li J B, Dundas I, Dong C G, Li G R, Trethowan R, Yang Z J, Hoxha S, Zhang P. 2020. Identification and characterization of a new stripe rust resistance gene Yr83 on rye chromosome 6R in wheat. Theoretical and Applied Genetics, 133, 1095–1107.

Li L H, Li X Q, Li P, Dong Y C, Zhao G S. 1997. Establishment of wheat–Agropyron cristatum alien addition lines I. Cytology of F₃, F₂BC₁, BC₄, and BC₃F₁ progenies. Acta Genetica Sinica, 24, 154–159. (in Chinese)

Li L H, Li X Q, Yang X M. 2006. Description Specification and Data Standard of Wheat Germplasm Resources. China Agricultural Publishing Society, Beijing. pp. 2–3. (in Chinese)

Li X L, Zhang M S, Ren M J, Lu X, Ji N, Wang X H, Xu R H. 2017. Molecular regulation mechanism of anthocyanin synthesis in purple wheat. Journal of Plant Physiology, 53, 521–530.

Lin Y D, Zhou S H, Liang X Z, Guo B J, Han B, Han H M, Zhang J P, Lu Y Q, Zhang Z, Yang X M, Li X Q, Liu W H, Li L H. 2022. Chromosomal mapping of a locus associated with adult-stage resistance to powdery mildew from Agropyron cristatum chromosome 6PL in wheat. Theoretical and Applied Genetics, 135, 2861–2873.

Lin Y D, Zhou S H, Liang X Z, Han B, Yang J L, Guo B J, Zhang J P, Han H M, Liu W H, Yang X M, Li X Q, Li L H. 2023. Introgression of chromosome 6PL terminal segment from Agropyron cristatum to increase both grain number and grain weight in wheat. The Crop Journal, 11, 878–886.

Liu L, Luo Q L, Li H W, Li B, Li Z S, Zheng Q. 2018. Physical mapping of the blue-grained gene from Thinopyrum ponticum chromosome 4Ag and development of blue-grain-related molecular markers and a FISH probe based on SLAF-seq technology. Theoretical and Applied Genetics, 131, 2359–2370.

Liu W, Koo D H, Friebe B, Gill B S. 2016. A set of Triticum aestivum–Aegilops speltoides Robertsonian translocation lines. Theoretical and Applied Genetics, 129, 2359–2368.

Liu X, Zhang M H, Jiang X M, Li H, Jia Z J, Hao M, Jiang B, Huang L, Ning S H, Yuan Z W, Chen X J, Chen X, Liu D C, Liu B L, Zhang LQ. 2021. TbMYC4A is a candidate gene controlling the blue aleurone trait in a wheat–Triticum boeoticum substitution line. Frontiers in Plant Science, 12, 762265.

Lv M J, Lu Y Q, Li H H, Pan C L, Guo Y, Zhang J P, Yang X M, Li Q C, Liu W H, Li L H. 2016. Transferring desirable genes from Agropyron cristatum 7P chromosome into common wheat. PLoS ONE, 11, e0159577.

Nomi Y, Iwasaki-Kurashige K, Matsumoto H. 2019. Therapeutic effects of anthocyanins for vision and eye health. Molecules, 24, 3311.

Ou J M, Li S R, Pang Q H, Tao J, Zhou Q, Du X Y. 2004. Genetic analysis of black seed coat in wheat Luozhen No. 1. Journal of Southwest Agricultural University, 26, 185–190. (in Chinese)

Padhy A K, Kaur P, Singh S, Kashyap L, Sharma A. 2024. Colored wheat and derived products: Key to global nutritional security. Critical Reviews in Food Science and Nutrition, 64, 1894–1910.

Qi K, Han H, Zhang J, Zhou S, Li X, Yang X, Liu W, Lu Y, Li L. 2021. Development and characterization of novel Triticum aestivum–Agropyron cristatum 6P Robertsonian translocation lines. Molecular Breeding, 41, 59.

Qian X K. 2019. Genetic regularity and gene mapping of purple seed character of guizhou purple wheat no.1. MSc thesis, Guizhou University, China. (in Chinese)

Quinlan A R. 2014. BEDTools: The swiss-army tool for genome feature analysis. Current Protocols in Bioinformatics, 47, 1–45.

Saini P, Kumar N, Kumar S, Mwaurah P W, Panghal A, Attkan A K, Singh V K, Garg M K, Singh V. 2020. Bioactive compounds, nutritional benefits and food applications of colored wheat: A comprehensive review. Critical Reviews in Food Science and Nutrition, 61, 3197–3210.

Sharma N, Tiwari V, Vats S, Kumari A, Chunduri V, Kaur S, Kapoor P, Garg M. 2020. Evaluation of anthocyanin content, antioxidant potential and antimicrobial activity of black, purple and blue colored wheat flour and wheat–grass juice against common human pathogens. Molecules, 25, 5785.

Shen X, Li G, Zhang L. 2013. Identification and mapping of the BaThb gene for the blue aleurone trait from Th. bessarabicum on chromosome 4J of wheat. Euphytica, 190, 451–460.

Song L Q, Li L J, Han H M, Gao A N, Yang X M, Li L H, Liu W H. 2013. Efficient induction of wheat–Agropyron cristatum 6P translocation lines and GISH detection. PLoS ONE, 8, e69501.

Song L Q, Lu YQ, Zhang J P, Pan C, Yang X M, Li X Q, Liu W H, Li L H. 2016. Cytological and molecular analysis of wheat–Agropyron cristatum translocation lines with 6P chromosome fragments conferring superior agronomic traits in common wheat. Genome, 59, 840–850.

Song X W, Wei J B, Di S K, Pang Y Z. 2019. Advances in the regulation of anthocyanin transcription factors and metabolic engineering. Journal of Plants, 54, 133–156.

Sun Q, Sun B Q, Wang J H. 2003. Genetic study on grain pigment of black wheat. Seed, 4, 33–34, 37.

Sun Y Y, Lu M J, Han H M, Zhou S H, Lu Y Q, Liu W H, Yang X M, Li X Q, Zhang J P, Liu X, Li L H. 2021. Identification and fine mapping of alien fragments associated with enhanced grain weight from Agropyron cristatum chromosome 7P in common wheat backgrounds. Theoretical and Applied Genetics, 134, 3759–3772.

Tang S Y, Tang Z X, Qiu L, Yang Z J, Li G R, Lang T, Zhu W Q, Zhang J H, Fu S L. 2018. Developing new oligo probes to distinguish specific chromosomal segments and the A, B, D genomes of wheat (Triticum aestivum L.) using ND-FISH. Frontiers in Plant Science, 9, 1104.

Tereshchenko O Y, Pshenichnikova T A, Salina E A, Khlestkina E K. 2012. Development and molecular characterization of a novel wheat genotype having purple seeds colour. Cereal Research Communications, 40, 210–214.

Tiwari V, Kamboj A, Sheoran B, Chaudhary E, Yadav M, Kumari A, Krishania M, Ali U, Tiwari A, Garg M, Bhatnagar A. 2025. Anthocyanin-rich black wheat as a functional food for managing type 2 diabetes mellitus: A study on high fat diet-streptozotocin-induced diabetic rats. Food & Function, 16, 3273–3295.

Tucker E J, Baumann U, Kouidri A, Suchecki R, Baes M, Garcia M, Okada T, Dong C, Wu Y, Sandhu A, Singh M, Langridge P, Wolters P, Albertsen M C, Cigan A M, Whitford R. 2017. Molecular identification of the wheat male fertility gene Ms1 and its prospects for hybrid breeding. Nature Communications, 8, 869.

Wang Q, Cao H M, Wang J C, Gu Z R, Lin Q Y, Zhang Z Y, Zhao X Y, Gao W, Zhu H J, Yan H B, Yan J J, Hao Q T, Zhang Y W. 2024. Fine-mapping and primary analysis of candidate genes associated with seed coat color in mung bean (Vigna radiata L.). Journal of Integrative Agriculture, 23, 2571–2588.

Wang X, Han B H, Sun Y Y, Kang X L, Zhang M, Han H M, Zhou S H, Liu W H, Lu Y Q, Yang X M, Li X Q, Zhang J P, Liu X, Li L H. 2022. Introgression of chromosome 1P from Agropyron cristatum reduces leaf size and plant height to improve the plant architecture of common wheat. Theoretical and Applied Genetics, 135, 1951–1963.

Wang Y, Yao X H, An L K, Yao Y H, Bai Y X, Wu K L. 2023. Research progress on the mechanism of grain color formation and gene mapping in cereals. Anhui Agricultural Sciences, 51, 1–6. (in Chinese)

Wickham H. 2016. ggplot2: Elegant graphics for data analysis. Journal of the Royal Statistical Society, 174, 245–246.

Wu J, Yang X, Wang H, Li H, Li L, Li X, Liu W. 2006. The introgression of chromosome 6P specifying for increased numbers of florets and kernels from Agropyron cristatum to wheat. Theoretical and Applied Genetics, 114, 13–20.

Xu P, Zhang Z B, Zhang J P, Li F R. 2022. Gene Discovery and germplasm resources breeding and utilization of color wheat. Journal of Plant Genetic Resources, 23, 1549–1571.

Xu S R, Jiang B, Han H M, Ji X J, Zhang J P, Zhou S H, Yang X M, Li X Q, Li L H, Liu W H. 2023. Genetic effects of Agropyron cristatum 2P chromosome translocation fragments in wheat background. Journal of Integrative Agriculture, 22, 52–62.

Yang G H, Yang X M, Wang R H, Gao A N, Li L H, Liu W H. 2010. The inhibiting effect of 1.4 recombinant P chromosome of wheat–Agropyron cristatum addition line on the Ph gene. Science Bulletin, 55, 153–157.

Ying K, Meng T Q, Gao W N, Yu Y H, Tian H Z, Wu J, Zhang Z M, Liu Y X. 2025. Comprehensive evaluation of main agronomic traits and nutritional quality of colored wheat germplasms. Journal of Triticeae Crops, 45, 337–348. (in Chinese)

Yu L L, Beta T. 2015. Identification and antioxidant properties of phenolic compounds during production of bread from purple wheat grains. Molecules, 20, 15525–15549.

Zeven A C. 1991. Wheats with purple and blue grains: A review. Euphytica, 56, 243–258.

Zhao X, Guo Y, Kang L, Yin C, Bi A, Xu D, Zhang Z, Zhang J, Yang X, Xu J, Xu S, Song X, Zhang M, Li Y, Kear P, Wang J, Liu Z, Fu X, Lu F. 2023. Population genomics unravels the Holocene history of bread wheat and its relatives. Nature Plants, 9, 403–419.

Zhang J, Zhang J P, Liu W H, Han H M, Lu Y Q, Yang X M, Li X Q, Li L H. 2015. Introgression of Agropyron cristatum 6P chromosome segment into common wheat for enhanced thousand-grain weight and spike length. Theoretical and Applied Genetics, 128, 1827–1837.

Zhang J, Zhang J P, Liu W H, Wu X Y, Yang X M, Li X Q, Lu Y Q, Li L H. 2016. An intercalary translocation from Agropyron cristatum 6P chromosome into common wheat confers enhanced kernel number per spike. Planta, 244, 853–864.

Zhang Q. 2021. Purple tomatoes, black rice and food security. Nature Reviews Genetics, 22, 414.

Zhang Z, Han H M, Liu W H, Song L Q, Zhang J P, Zhou S H, Yang X M, Li X Q, Li L H. 2019. Deletion mapping and verification of an enhanced-grain number per spike locus from the 6PL chromosome arm of Agropyron cristatum in common wheat. Theoretical and Applied Genetics, 132, 2815–2827.

Zhang Z, Song L Q, Han H M, Zhou S H, Zhang J P, Yang X M, Li X Q, Liu W H, Li L H. 2017. Physical localization of a locus from Agropyron cristatum conferring resistance to stripe rust in common wheat. International Journal of Molecular Sciences, 18, 2403.

Zhang Z, Zhou S H, Liu W H, Song L Q, Zhang J P, Han H M, Yang X M, Lin Y D, Li Q C, Li L H. 2021. Molecular cytogenetic analysis of the introgression between Agropyron cristatum P genome and wheat genome. International Journal of Molecular Sciences, 22, 11208.

Zhang Z B, Xu P, Li F L, Zhang W S, Li F R. 2024. Correlation of agronomic traits and micronutrient elements of colored wheat. Chinese Journal of Eco-Agriculture, 32, 1669–1678.

Zheng Q, Li B, Zhang X Y, Mu S M, Zhou H P, Li Z S. 2006. Physical mapping of the blue-grained gene from Thinopyrum ponticum by GISH and FISH in a set of translocation lines with different seed colors in wheat. Genome, 49, 1109–1114.

Zhu T, Wang L, Rimbert H, Rodriguez J C, Deal K R, De Oliveira R, Choulet F, Keeble-Gagnère G, Tibbits J, Rogers J, Eversole K, Appels R, Gu Y Q, Mascher M, Dvorak J, Luo M C. 2021. Optical maps refine the bread wheat Triticum aestivum cv. Chinese Spring genome assembly. The Plant Journal, 107, 303–314.

Zhu Z X, Zhang Y L, Jia J, Li W L, Zhao L D, Meng F G, Gai H M, Xu X X, Zhao C X. 2023. Analysis of dry matter accumulation and yield formation of different color wheat varieties (lines). Journal of North China Agriculture, 38, 128–138. (in Chinese)

Zhou S H, Zhang J P, Che Y H, Liu W H, Lu Y Q, Yang X M, Li Q C, Jia J Z, Liu X, Li L H. 2018. Construction of Agropyron Gaertn. genetic linkage maps using a wheat 660K SNP array reveals a homoeologous relationship with the wheat genome. Plant Biotechnology Journal, 16, 818–827.

Zong X F. 2006. Study on the growth characteristics, grain nutrient and antioxidation of black wheat (T. aestivum L.). Ph D thesis, Southwest University, China. (in Chinese)

Zong X F, Yan R, Li B X, Wang S G. 2017. Dynamic changes of pigments and related enzymes in blue- and purple-grained wheat kernels. Journal of Southwest University (Natural Science Edition), 39, 1–7. (in Chinese)

Zong Y, Li G, Xi X, Sun X, Li S, Cao D, Zhang H, Liu B. 2019. A bHLH transcription factor TsMYC2 is associated with the blue grain character in triticale (Triticum × Secale). Plant Cell Reports, 38, 1291–1298.