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Kernel softness in wheat is determined by starch granule bound Puroindoline proteins

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Abstract

Wheat has a vital position in agriculture because it is a staple food for masses and variation in grain hardness governs its applications. Soft wheats have softer endosperm texture that mills easily, so needs less energy to mill, produces smaller particles, and small amount of starch is damaged after milling as compared to hard wheat. Soft texture results from higher level of friabilin whereas hard texture results from low level of friabilin on starch granule surface. Friabilin, a marker of kernel texture is primarily composed of Puroindolines (PINs) and its genes (Pins) are located on the Hardness (Ha) locus. The Pins are the molecular-genetic basis of kernel softness in wheat. When both Pins are in their ‘wild state’ (Pina-D1a and Pinb-D1a), wheat kernel is soft. Absence or mutation in one of the Pins results in hard grain texture with different effects on end use and milling qualities. Pina-D1b genotypes gave harder grain texture, higher protein content, water absorption of flour, damaged starch granules and greater flour yield than hard wheat. Recently, other Pins like genes, Pin b variant genes located on the long arm of chromosome 7A were reported in bread wheat with more than 70% similarity to Pinb (Pinb-D1a) at the DNA level. Other genes located on chromosomes 1A, 2A, 5A, 7A, 5B, 2D and 6D also affect kernel texture. However the main determinants are the variants in the allelic diversity of Puroindoline family genes. Contemporary studies show that Pins are multifunctional family of genes having a range of functions from grain hardness to natural defense against insects and pathogens such as viruses, bacteria and fungi.

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References

  • Ali I, Sardar Z, Rasheed A, Mahmood T (2015) Molecular characterization of the Puroindoline-a and b alleles in synthetic hexaploid wheats and in silico functional and structural insights into Pina-D1. J Theor Biol 376:1–7

    Article  CAS  PubMed  Google Scholar 

  • Andrushchenko VV, Aarabi MH, Nguyen LT, Prenner EJ, Vogel HJ (2008) Thermodynamics of the interactions of tryptophan-rich cathelicidin antimicrobial peptides with model and natural membranes. Biochim Biophys Acta Biomembr 1778:1004–1014

    Article  CAS  Google Scholar 

  • Baker RJ, Dyck PL (1975) Relation of several quality characteristics to hardness in two spring wheat crosses. Can J Plant Sci 55:625–627

    Article  Google Scholar 

  • Beecher B, Bettge A, Smidansky E, Giroux MJ (2002) Expression of wild-type pinB sequence in transgenic wheat complements a hard phenotype. Theor Appl Genet 105:870–877

    Article  CAS  PubMed  Google Scholar 

  • Bhave M, Morris CF (2008a) Molecular genetics of puroindolines and related genes: allelic diversity in wheat and other grasses. Plant Mol Biol 66:205–219

    Article  CAS  PubMed  Google Scholar 

  • Bhave M, Morris CF (2008b) Molecular genetics of puroindolines and related genes: regulation of expression, membrane binding properties and applications. Plant Mol Biol 66:221–231

    Article  CAS  PubMed  Google Scholar 

  • Bibi A, Ajmal S, Rasheed A, Gul-Kazi A, Mujeeb-Kazi A (2012) High molecular weight (HMW) glutenin subunit compositions of the Elite-II synthetic hexaploid wheat sub-set (Triticum turgidum × Aegilops tauschii; 2n = 6x = 42; AABBDD). Plant Genet Resour 10:1–4

    Article  CAS  Google Scholar 

  • Blochet JE, Kaboulou A, Compoint JP, Marion D (1991) Amphiphilic proteins from wheat flour: specific extraction, structure and lipid binding properties. In: Bushuk W, Tkachuk R (eds) Gluten proteins 1990. American Association of Cereal Chemists, Saint Paul Minnesota, pp 314–325

    Google Scholar 

  • Blochet JE, Chevalier C, Forest E, Pebay-Peyroula E, Gautier MF, Joudrier P, Pezolet M, Marion D (1993) Complete amino acid sequence of puroindoline, a new basic and cystine-rich protein with a unique tryptophan-rich domain, isolated from wheat endosperm by Triton X-114 phase partitioning. FEBS Lett 329:336–340

    Article  CAS  PubMed  Google Scholar 

  • Bordier C (1981) Phase separation of integral membrane proteins in Triton X-114 solution. J Biol Chem 256:1604–1607

    CAS  PubMed  Google Scholar 

  • Cane K, Sharp PJ, Eagles HA, Eastwood RF, Hollamby GJ, Kuchel K, Lu M, Martin PJ (2008) The effects on grain quality traits of a grain serpin protein and the VPM1 segment in southern Australian wheat breeding. Aust J Agric Res 59:883–895

    Article  CAS  Google Scholar 

  • Chang C, Zhang HP, Xu J, Li WH, Liu GT, You MS, Li BY (2006) Identification of allelic variations of puroindoline genes controlling grain hardness in wheat using a modified denaturing PAGE. Euphytica 152:225–234

    Article  CAS  Google Scholar 

  • Chantret N, Salse J, Sabot F, Rahman S, Bellec A, Laubin B, Dubois I, Dossat C, Sourdille P, Joudrier P, Gautier MF, Cattolico L, Beckert M, Aubourg S, Weissenbach J, Caboche M, Bernard M, Leroy P, Chalhoub B (2005) Molecular basis of evolutionary events that shaped the hardness locus in diploid and polyploid wheat species Triticum and Aegilops. Plant Cell 17:1033–1045

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  • Charnet P, Molle G, Marion D, Rousset M, Lullien-Pellerin V (2003) Puroindolines form ion channels in biological membranes. Biophys J 84:2416–2426

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  • Chen F, He Z, Xia X, Lillemo M, Morris C (2005) A new puroindoline-b mutation present in Chinese winter wheat cultivar Jingdong 11. J Cereal Sci 42:267–269

    Article  CAS  Google Scholar 

  • Chen F, Xia XC, Wang DS, Lillemo M, He ZH (2006) Detection of allelic variation for Puroindoline alleles in CIMMYT germplasm developed from synthetic wheat crossing with common wheats. Sci Agric Sin 39:440–447

    CAS  Google Scholar 

  • Chen F, He ZH, Chen DS, Zhang CL, Zhang Y, Xia XH (2007) Influence of puroindoline alleles on milling performance and qualities of Chinese noodles, steamed bread and pan bread in spring wheats. J Cereal Sci 45:59–66

    Article  CAS  Google Scholar 

  • Chen F, Zhang F, Morris CF, He ZH, Xia XC, Cui D (2010) Molecular characterization of the Puroindolinea-D1b allele and development of an STS marker in wheat (Triticumaestivum L.). J Cereal Sci 52:80–82

    Article  CAS  Google Scholar 

  • Chen F, Li H, Cui D (2013) Discovery, distribution and diversity of Puroindoline-D1 genes in bread wheat from five countries (Triticum aestivum L.). BMC Plant Biol 13:125. doi:10.1186/1471-2229-13-125

    Article  PubMed  PubMed Central  Google Scholar 

  • Clarke B, Rahman S (2005) A microarray analysis of wheat grain hardness. Theor Appl Genet 110:1259–1267

    Article  CAS  PubMed  Google Scholar 

  • Corona V, Gazza L, Boggini G, Pogna NE (2001) Variation in friabilin composition as determined by A-PAGE fractionation and PCR amplification, and its relationship to grain hardness in bread wheat. J Cereal Sci 34:243–250

    Article  CAS  Google Scholar 

  • Day L, Bhandari DG, Greenwell P, Leonard SA, Schofield JD (2006) Characterization of wheat puroindoline proteins. FEBS J 273:5358–5373

    Article  CAS  PubMed  Google Scholar 

  • Digeon JF, Guiderdoni E, Alary R, Michaux-Ferrière N, Joudrier P, Gautier MF (1999) Cloning of a wheat puroindoline gene promoter by IPCR and analysis of promoter regions required for tissue-specific expression in transgenic rice seeds. Plant Mol Biol 39:1101–1112

    Article  CAS  PubMed  Google Scholar 

  • Dubreil L, Gaborit T, Bouchet B, Gallant DJ, Broekaert W, Quillien L, Marion D (1998) Spatial and temporal distribution of the major isoforms of puroindolines (puroindoline-a and puroindoline-b) and non-specific lipid transfer protein (ns-LTPe1) of Triticum aestivum seeds: relationships with their in vitro antifungal properties. Plant Sci 138:121–135

    Article  CAS  Google Scholar 

  • Eagles HA, Cane K, Eastwood RF, Hollamby GJ, Kuchel H, Martin PJ, Cornish GB (2006) Contributions of glutenin and puroindoline genes to grain quality traits in southern Australian wheat breeding programs. Aust J Agric Res 57:179–186

    Article  CAS  Google Scholar 

  • Edwards MA, Osborne BG, Henry RJ (2010) Puroindoline genotype, starch granule size distribution and milling quality of wheat. J Cereal Sci 52:314–320

    Article  CAS  Google Scholar 

  • Encinas R (2012) Report from the field. Wheat: a rich man’s crop. Conservation Agriculture Events, CIMMYT

  • Evrard A, Lagarde V, Joudrier P, Gautier M (2008) Puroindoline-a and puroindoline-b interact with the Saccharomyces cerevisiae plasma membrane through different amino acids present in their tryptophan-rich domain. J Cereal Sci 48:379–386

    Article  CAS  Google Scholar 

  • Galande AA, Tiwari R, Ammiraju JSS, Santra DK, Lagu MD, Rao VS, Gupta VS, Misra BK, Nagarajan S, Ranjekar PK (2001) Genetic analysis of kernel hardness in bread wheat using PCRbased markers. Theor Appl Genet 103:601–606

    Article  CAS  Google Scholar 

  • Gasparis S, Orczyk W, Nadolska-Orczyk A (2013) Sina and Sinb genes in triticale do not determine grain hardness contrary to their orthologs Pina and Pinb in wheat. BMC Plant Biol 13:190. doi:10.1186/1471-2229-13-190

    Article  PubMed  PubMed Central  Google Scholar 

  • Gautier MF, Aleman ME, Guirao A, Marion D, Joudrier P (1994) Triticum aestivum puroindolines, two basic cysteine-rich seed proteins: cDNA sequence analysis and developmental gene expression. Plant Mol Biol 25:43–57

    Article  CAS  PubMed  Google Scholar 

  • Gautier MF, Cosson P, Guirao A, Alary R, Joudrier P (2000) Puroindoline genes are highly conserved in diploid ancestor wheats and related species but absent in tetraploid Triticum species. Plant Sci 153:81–91

    Article  CAS  Google Scholar 

  • Gazza L, Niglio A, Vaccino P, Pogna NE (2003) The long arm of chromosome 5D of bread wheat contains a Pina-D1a-like sequence. In: Pogna NE, Romano M, Pogna E, Galterio G (eds) Proceedings of the 10th international wheat genetics symposium, vol 3. Paestum, Italy, pp 1330–1332

    Google Scholar 

  • Gazza L, Nocente F, Ng PKW, Pogna NE (2005) Genetic and biochemical analysis of common wheat cultivars lacking puroindoline a. Theor Appl Genet 110:470–478

    Article  CAS  PubMed  Google Scholar 

  • Gedye KR, Morris CF, Bettge AD (2004) Determination and evaluation of the sequence and textural effects of the puroindoline a and puroindoline b genes in a population of synthetic hexaploid wheat. Theor Appl Genet 109:1597–1603

    Article  CAS  PubMed  Google Scholar 

  • Geng H, Beecher BS, He Z, Kiszonas AM, Morris CF (2012) Prevalence of Puroindoline D1 and Puroindoline b-2 variants in U.S. Pacific Northwest wheat breeding germplasm pools, and their association with kernel texture. Theor Appl Genet 124:1259–1269

    Article  CAS  PubMed  Google Scholar 

  • Giroux MJ, Morris CF (1997) A glycine to serine change in puroindoline b is associated with wheat grain hardness and low levels of starch-surface friabilin. Theor Appl Genet 95(5–6):857–864

    Article  CAS  Google Scholar 

  • Giroux MJ, Morris CF (1998) Wheat grain hardness results from highly conserved mutations in the friablin components puroindoline a and b. Proc Natl Acad Sci USA 95:6262–6266

    Article  CAS  PubMed  PubMed Central  Google Scholar 

  • Giroux MJ, Talbert L, Habernicht DK, Lanning S, Hemphill A, Martin JM (2000) Association of puroindoline sequence type and grain hardness in hard red spring wheat. Crop Sci 30:370–374

    Article  Google Scholar 

  • Gollan P, Smith K, Bhave M (2007) Gsp-1 genes comprise a multigene family in wheat that exhibits a unique combination of sequence diversity yet conservation. J Cereal Sci 45:184–198

    Article  CAS  Google Scholar 

  • Greenwell P (1987) Wheat starch granule proteins and their technological significance. In: Murray L (eds) Proceedings of the 37th Australian cereal chemistry conference, 1 (Melbourne, Australia, 9–22 October 1987), Cereal Chemistry Division, Royal Australian Chemical Institute, Parkville, VC, Australia, pp 100–103

  • Greenwell P, Schofield JD (1986) A starch granule protein associated with endosperm softness in wheat. Cereal Chem 63:379–380

    CAS  Google Scholar 

  • Greenwell P, Schofield JD (1989) The chemical basis of grain hardness and softness. In: Salovaara H (eds) Wheat end-use properties. In: Proceedings ICC 89 symposium, University of Helsinki Lahti Research and Training Centre, Lahti, Finland, pp 59–72

  • Gupta RB, Shepherd KW (1990) Two-step one-dimensional SDS-PAGE analysis of LMW subunits of glutelin. I. Variation and genetic control of subunits in hexaploid wheats. Theor Appl Genet 80:65–74

    Article  CAS  PubMed  Google Scholar 

  • Hogg AC, Stripo T, Beecher B, Martin JM, Giroux MJ (2004) Wheat puroindolines interact to form friabilin and control wheat grain hardness. Theor Appl Genet 108:1089–1097

    Article  CAS  PubMed  Google Scholar 

  • Ikeda TM, Ohnishi N, Nagamine T, Oda S, Hisatomi T, Yano H (2005) Identification of new puroindoline genotypes and their relationship to flour texture among wheat cultivars. J Cereal Sci 41:1–6

    Article  CAS  Google Scholar 

  • Ikeda TM, Cong H, Suzuki T, Takata K (2010) Identification of new Pina null mutations among Asian common wheat cultivars. J Cereal Sci 51:235–237

    Article  CAS  Google Scholar 

  • International Wheat Genome Sequencing Consortium (2014) A chromosome-based draft sequence of the hexaploid bread wheat (Triticum aestivum) genome. Science 345:1251788. doi:10.1126/science.1251788

    Article  Google Scholar 

  • Jolly CJ, Rahman S, Korrt AA, Higgins TJV (1993) Characterization of the wheat Mr 15000 ‘grain softness protein’ and analysis of the relationship between its accumulation in the whole seed and grain softness. Theor Appl Genet 86:589–597

    Article  CAS  PubMed  Google Scholar 

  • Kumar R, Arora S, Singh K, Garg M (2015) Puroindoline allelic diversity in Indian wheat germplasm and identification of new allelic variants. Breed Sci 65(4):319–326. doi:10.1270/jsbbs.65.319

    Article  PubMed  PubMed Central  Google Scholar 

  • Law CN, Young CF, Brown JWS, Snape JW, Worland JW (1978) The study of grain protein control in wheat using whole chromosome substitution lines. In: Seed protein improvement by nuclear techniques. International Atomic Agency, Vienna, Austria, pp 483–502

  • Li G, He Z, Lillemo M, Sun Q, Xia X (2008) Molecular characterization of allelic variations at Pina and Pinb loci in Shandong wheat landraces, historical and current cultivars. J Cereal Sci 47:510–517

    Article  CAS  Google Scholar 

  • Lillemo M, Morris CF (2000) A leucine to proline mutation in puroindoline b is frequently present in hard wheats from Northern Europe. Theor Appl Genet 100:1100–1107

    Article  CAS  Google Scholar 

  • Lillemo M, Simeone MC, Morris CF (2002) Analysis of Puroindoline a and b sequences from Triticum aestivum cv. ‘Penawawa’ and related taxa. Euphytica 126:321–331

    Article  CAS  Google Scholar 

  • Lillemo M, Chen F, Xia XC, William M, Pena RJ, Trethowan R, He ZH (2006) Puroindoline grain hardness alleles in CIMMYT bread wheat. J Cereal Sci 44:86–92

    Article  CAS  Google Scholar 

  • Luo L, Zhang JR, Yang GX, Li Y, Li KX, He GY (2008) Expression of puroindoline a enhances leaf rust resistance in transgenic tetraploid wheat. Mol Biol Rep 35:195–200

    Article  CAS  PubMed  Google Scholar 

  • Martin CR, Rousser R, Brabec DL (1993) Development of a single-kernel wheat characterization system. Trans Am Soc Agric Eng 36:1399–1404

    Article  Google Scholar 

  • Martin JM, Frohberg RC, Morris CF, Talbert LE, Giroux MJ (2001) Milling and bread baking traits associated with puroindoline sequence type in hard red spring wheat. Crop Sci 41:228–234

    Article  CAS  Google Scholar 

  • Massa AN, Morris CF, Gill BS (2004) Sequence diversity of puroindoline-a, puroindoline-b, and the grain softness protein genes in Aegilops tauschii cross. Crop Sci 44:1808–1816

    Article  CAS  Google Scholar 

  • McIntosh RA, Devos KM, Dubcovsky WJ, Rogers WJ, Morris CF, Appels R, Anderson OA (2006) Catalogue of gene symbols for wheat: 2006 supplement. http://www.grain.jouy.inra.fr/ggpages/wgc/2006upd.html

  • Morris CF (2002) Puroindolines: the molecular genetic basis of wheat grain hardness. Plant Mol Biol 48:633–647

    Article  CAS  PubMed  Google Scholar 

  • Morris CF, Greenblatt GA, Bettge AD, Malkawi HI (1994) Isolation and characterisation of multiple forms of friabilin. J Cereal Sci 21:167–174

    Article  Google Scholar 

  • Morris CF, DeMacon VL, Giroux MJ (1999) Wheat grain hardness among chromosome 5D homozygous recombinant substitution lines using different methods of measurement. Cereal Chem 76:249–254

    Article  CAS  Google Scholar 

  • Morris CF, Lillemo M, Simeone MC, Giroux MJ, Babb SL, Kidwell KK (2001) Prevalence of puroindoline grain hardness genotypes among historically significant North American spring and winter wheats. Crop Sci 41:218–228

    Article  CAS  Google Scholar 

  • Mujeeb-Kazi A, Rosas V, Roldan S (1996) Conservation of the genetic variation of Triticum tauschii (Coss.) Schmalh. (Aegilops squarrosa auct. Non L.) in synthetic hexaploid wheats (T. turgidum L. × T. tauschii; 2n = 6x = 42, AABBDD) and its potential utilization for wheat improvement. Genet Resour Crop Evol 43:129–134

    Article  Google Scholar 

  • Norris KH, Hruschka WR, Bean MM, Slaughter DC (1989) A definition of wheat hardness using the near infrared reflectance spectroscopy. Cereal Food World 34:696–705

    Google Scholar 

  • Palumbo D, Iannaccone M, Porta A, Capparelli R (2010) Experimental antibacterial therapy with puroindolines, lactoferrin and lysozyme in Listeria monocytogenes-infected mice. Microb Infect 12:538–545

    Article  CAS  Google Scholar 

  • Pan Z, Song W, Meng F, Xu L, Liu B, Zhu J (2004) Characterization of genes encoding wheat grain hardness from Chinese cultivar Gaocheng 8901. Cereal Chem 82:38–43

    Google Scholar 

  • Peng J, Sun D, Nevo E (2011) Wild emmer wheat, Triticum dicoccoides, occupies a pivotal position in wheat domestication process. Aust J Crop Sci 5:1127–1143

    Google Scholar 

  • Pogna NE, Gazza L, Corona V, Zanier R, Niglio A, Mei E, Palumbo M, Boggini G (2002) Puroindolines and kernel hardness in wheat species. In: Ng PKW, Wrigley CW (eds) Wheat quality elucidation. AACC, St.Paul, pp 155–169

    Google Scholar 

  • Ram S, Jain N, Shoran J, Singh R (2005) New frame shift mutation in Puroindoline-b in Indian wheat cultivars Hyb65 and NI5439. J Plant Biochem Biotechnol 14:45–48

    Article  CAS  Google Scholar 

  • Ramalingam A, Palombo EA, Bhave M (2012) The Pinb-2 genes in wheat comprise a multigene family with great sequence diversity and important variants. J Cereal Sci 56:171–180

    Article  CAS  Google Scholar 

  • Rasheed A, Mahmood A, Gul-Kazi A, Ghafoor A, Mujeeb-Kazi A (2012) Allelic variation and composition of HMW-GS in advanced derivaties from D-genome synthetic hexaploid × Bread wheat cross combinations. Journal of Crop Science and Biotechnology 15:1–7

    Article  Google Scholar 

  • Reynolds NP, Martin JM, Giroux MJ (2010) Increased wheat grain hardness conferred by novel puroindoline haplotypes from Aegilops tauschii. Crop Sci 50:1718–1727

    Article  CAS  Google Scholar 

  • Riley R, Chapman V (1958) Genetic control of the cytologically diploid behaviour of hexaploid wheat. Nature 182:713–715

    Article  Google Scholar 

  • Rosicka-Kaczmarek J, Stasiuk M, Nebesny E, Komisarczyk A (2015) Fluorimetric studies of the interactions of wheat puroindolines with polar lipids on the surface starch granules. J Cereal Sci 66:53–58

    Article  CAS  Google Scholar 

  • Schofield JD, Greenwell P (1987) Wheat starch granule proteins and their technological significance. In: Morton ID (ed) Cereals in a European Context. Ellis Horwood, Chichester, pp 407–420

    Google Scholar 

  • Shewry PR, Jenkins J, Beaudoin F, Mills ENC (2004) The classification, functions and evolutionary relationships of plant proteins in relation to food allergens. In: Mills ENC, Shewry PR (eds) Plant food allergens. Blackwell Science, Oxford, pp 24–41

    Google Scholar 

  • Tanaka H, Morris CF, Haruna M, Tsujimoto H (2008) Prevalence of puroindoline alleles in wheat from eastern Asia including discovery of a new SNP in puroindoline b. Plant Genet Resour 6:142–152

    Article  CAS  Google Scholar 

  • Tranquilli G, Lijavetzky D, Muzzi G, Dubcovsky J (1999) Genetic and physical characterization of grain texture-related loci in diploid wheat. Mol Gen Genet 262:846–850

    Article  CAS  PubMed  Google Scholar 

  • Tranquilli G, Heaton J, Chicaiza O, Dubcovsky J (2002) Substitutions and deletions of genes related to grain hardness in wheat and their effect on grain texture. Crop Sci 42:1812–1817

    Article  CAS  Google Scholar 

  • Turnbull KM, Rahman S (2002) Endosperm texture in wheat. J Cereal Sci 36:327–337

    Article  CAS  Google Scholar 

  • Turnbull KM, Turner M, Mukai Y, Yamamoto M, Morell MK, Appels R, Rahman H (2003) The organization of genes tightly linked to the Ha locus in Aegilops tauschii, the Dgenome donor to wheat. Genome 46:330–338

    Article  CAS  PubMed  Google Scholar 

  • Vakhitov VA, Chemeris AV, Sabirzhanov BE, Akhunov ED, Kulikov AM, Nikonorov IM, Gimalov FR, Bikbulatova SM, Baĭmiev AK (2003) Phylogeny of Triticum L. and Aegilops L. genuses inferred from a comparative analysis of nucleotide sequences in promoter rDNA regions of individual species. Genetika 39:5–17

    CAS  PubMed  Google Scholar 

  • Wilkinson M, Wan Y, Tosi P, Leverington M, Snape J, Mitchel RAC, Shewry PR (2008) Identification and genetic mapping of variant forms of puroindoline b expressed in developing wheat grain. J Cereal Sci 48:722–728

    Article  CAS  Google Scholar 

  • Worzella WW, Cutler GH (1939) A critical study of techniques for measuring granulation in wheat meal. J Agric Res 58:329–341

    Google Scholar 

  • Xia L, Chen F, He Z, Chen X, Morris CF (2005) Occurrence of puroindoline alleles in Chinese winter wheats. Cereal Chem 82:38–43

    Article  CAS  Google Scholar 

  • Yang W, Liu D, Li J, Zhang L, Wei H, Hu X, Zheng Y, He Z, Zou Y (2009) Synthetic hexaploid wheat and its utilization for wheat genetic improvement in China. J Genet Genom 36:539–546

    Article  CAS  Google Scholar 

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Iftikhar, A., Ali, I. Kernel softness in wheat is determined by starch granule bound Puroindoline proteins. J. Plant Biochem. Biotechnol. 26, 247–262 (2017). https://doi.org/10.1007/s13562-016-0387-1

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