Journal of Applied Genetics

, Volume 50, Issue 3, pp 185–197 | Cite as

The determinants of grain texture in cereals

  • A. Nadolska-Orczyk
  • S. Gasparis
  • W. Orczyk
Review Article


Kernel hardness is an important agronomic trait that influences end-product properties. In wheat cultivars, this trait is determined by thePuroindoline a (Pina) andPuroindoline b (Pinb) genes, located in theHardness locus (Ha) on chromosome 5DS of the D genome. Wild type alleles code puroindoline a (PINA) and puroindoline b (PINB) proteins, which form a 15-kDa friabilin present on the surface of water-washed starch granules. Both the proteins are accumulated in the starch endosperm cells and aleurone of the mature kernels.Puroindoline-like genes coding puroindoline-like proteins in the starch endosperm occur in some of the genomes of Triticeae and Aveneae cereals. Orthologs are present in barley, rye and oats. However, some genomes of these diploid and polyploid cereals, like that ofTriticum turgidum var.durum (AABB) lack thepuroindoline genes, having a very hard kernel texture. The two wild type alleles in opposition (dominant loci) control the soft pheno-type. Mutation either inPina orPinb or in both leads to a medium-hard or hard kernel texture. The most frequent types ofPin mutations are point mutations within the coding sequence resulting in the substitution of a single amino acid or a null allele. The latter is the result of a frame shift determined by base deletion or insertion or a one-point mutation to the stop codon. The lipid-binding properties of the puroindolines affect not only the dough quality but also the plants’ resistance to pathogens. Genetic modification of cereals withPuroindoline genes and/or their promoters enable more detailed functional analyses and the production of plants with the desired characteristics.


grain hardness puroindoline proteins Pin mutations puroindoline gene modifications Triticum sp 


Unable to display preview. Download preview PDF.

Unable to display preview. Download preview PDF.


  1. Amoroso MG, Longobardo L, Capprarelli R, 2004. Real time RT-PCR and flow cytometry to investigate wheat kernel hardness: role of puroindoline genes and proteins. Biotechnol Lett 26: 1731–1737.CrossRefPubMedGoogle Scholar
  2. Baker RJ, 1977. Inheritance of kernel hardness in spring wheat. Crop Sci 17: 960–962.CrossRefGoogle Scholar
  3. Bhave M., Morris, C. F., 2008a. Molecular genetics of puroindolines and related genes: allelic diversity in wheat and other grasses. Plant Mol Biol 66, 205–219.CrossRefPubMedGoogle Scholar
  4. 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.CrossRefPubMedGoogle Scholar
  5. Beecher B, Smidansky ED, See D, Blake TK, Giroux MJ, 2001. Mapping and sequence analysis of barley hordoindolines. Theor Appl Genet 102: 833–840.CrossRefGoogle Scholar
  6. Beecher B, Bettge A, Smidansky E, Giroux MJ, 2002. Expression of wild-typePinB sequence in transgenic wheat complements a hard phenotype. Theor Appl Genet 105: 870–877.CrossRefPubMedGoogle Scholar
  7. Blochet J-E, Chevalier C, Forest E, Pebay-Peyroula E, Gautier MF, Joudrier P, et al. 1993. Complete amino acid sequence of puroindoline, a new basic and cysteine-rich protein with a unique tryptophan-rich domain, isolated from wheat endosperm by Triton X-114 phase partitioning. FEBS Letters 329: 336–340.CrossRefPubMedGoogle Scholar
  8. Breseghello F, Finney PL, Gaines C, Andrews L, Tanaka J, Penner G, Sorrells ME, 2005. Genetic loci related to kernel quality differences between a soft and a hard wheat cultivar. Crop Sci 45: 1685–1695.CrossRefGoogle Scholar
  9. Broekaert WF, Marin W, Terras FRG, De Bolle MFC, Thevissen K, De Samblanx GW, Osborn RW, 1997. Antimicrobial peptides from plants. Crit Rev Plant Sci 16: 297–323.Google Scholar
  10. Bushuk W, 1998. Wheat breeding for end-product use. Euphytica 100: 137–145.CrossRefGoogle Scholar
  11. Campbell KG, Bergman CJ, Gaulberto DG, Anderson JA, Giroux MJ, Hareland G, Fulcher RG, et al. 1999. Quantitative trait loci associated with kernel traits in a soft × hard wheat cross. Crop Sci 39: 1184–1195.CrossRefGoogle Scholar
  12. Caldwell KS, Langridge P, Powell W, 2004. Comparative sequence analysis of the region harboring the hardness locus in barley and its colinear region in rice. Plant Physiol 136: 1–14.CrossRefGoogle Scholar
  13. Cane K, Spackman M, Eagles HA, 2004. Puroindoline genes and their effects on grain quality traits in southern Australian wheat cultivars. Aust J Agric Res 55: 89–95CrossRefGoogle Scholar
  14. Capparelli R, Borriello G, Giroux MJ, Amoroso MG, 2003. Puroindoline A-gene expression is involved in association of puroindolines to starch. Theor Appl Genet 107: 1463–1468.CrossRefPubMedGoogle Scholar
  15. Capparelli R, Amoroso MG, Palumbo D, Iannaccone M, Faleri C, Cresti M, 2005. Two plant puroindolines colocalize in wheat seed andin vitro synergistically fight against pathogens. Plant Mol Biol 58: 857–867.CrossRefPubMedGoogle Scholar
  16. Chang C, Zhang H, Xu J, Li W, Liu G, You M, Li B, 2006. Identification of allelic variations of puroindoline genes controlling grain hardness in wheat using a modified denaturing PAGE. Euphytica 152: 255–234.CrossRefGoogle Scholar
  17. Chantret N, Cenci A, Sabot F, Anderson O, Dubcovsky J, 2004. Sequencing of theTriticum monococcum Hardness locus reveals good microcolinearity with rice. Mol Genet Genomics 271: 377–386.CrossRefPubMedGoogle Scholar
  18. Chantret N, Salse J, Sabot F, Rahman S, Bellec A, Laubin B, et al. 2005. Molecular basis of evolutionary events that shaped theHardness locus in diploid and polyploid wheat species (Triticum andAegilops). Plant Cell 17: 1033–1045.CrossRefPubMedGoogle Scholar
  19. Chen F, He ZH, Xia XC, Lillemo M, Morris CF, 2005. A new puroindoline b mutation present in Chinese winter wheat cultivar Jingdong 11. J Cereal Sci 42: 267–269.CrossRefGoogle Scholar
  20. Chen F, He ZH, Xia XC, Xia LQ, Zhang XY, Lillemo M, Morris CF, 2006. Molecular and biochemical characterization of puroindoline a and b alleles in Chinese landraces and historical cultivars. Theor Appl Genet 112: 400–409.CrossRefPubMedGoogle Scholar
  21. Chen F, Yu Y, Xia X, He Z, 2007. Prevalence of a novel puroindoline b allele in Yunnan endemic wheats (Triticum aestivum ssp.Yunnanense King). Euphytica 156: 39–46.CrossRefGoogle Scholar
  22. Clarke B, Rahman S, 2005. A microarray analysis of wheat grain hardness. Theor Appl Genet 110: 1259–1267.CrossRefPubMedGoogle Scholar
  23. Corona V, Gazza L, Zanier R, Pogna NE, 2001a. A tryptophan-to-arginine change in the tryptophanrich domain of puroindoline b in five French bread wheat cultivars (Triticum aestivum L.). J Genet Breed 55: 187–189.Google Scholar
  24. Corona V, Gazza L, Boggini G, Pogna NE, 2001b. 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.CrossRefGoogle Scholar
  25. Darlington HF, Rouster J, Hoffman L, Halford NG, Shewry PR, Simpson DJ, 2001. Identyfication and molecular characterization of hordoindolines from barley grain. Plant Mol Biol 47: 785–794.CrossRefPubMedGoogle Scholar
  26. Darlington HF, Tecsi L, Harris N, Griggs DL, Cantrell IC, Shewry PR, 2000. Starch granule associated proteins in barley and wheat. J Cereal Sci 32: 21–29.CrossRefGoogle Scholar
  27. Digeon J-F, Guiderdoni E, Alary R, Michaux-Ferričre N, Joudrier P, Gautier M-F, 1999. Cloning of a wheat puroindoline gene promoter by IPCR and analysis of promoter regions requiredfortissue-specific expression in transgenic rice seeds. Plant Mol Biol 39: 1101–1112.CrossRefPubMedGoogle Scholar
  28. Douliez J-P, Michon T, Elmorjani K, Marion D, 2000. Structure, biological and technological functions of lipid transfer proteins and indolines, the major lipid binding proteins from cereal kernels. J Cereal Sci 32: 1–20.CrossRefGoogle Scholar
  29. Dubreil L, Compoint JP, Marion D, 1997. Interaction of puroindoline with wheat flour polar lipids determines their foaming properties. J Agric Food Chem 45: 108–116.CrossRefGoogle Scholar
  30. Dubreil L, Gaborit T, Bouchet B, Gallant DJ, Broekaert WF, Quillien L, Marion D, 1998a. Spatial and temporal distribution of the major isoforms of puroindolines (puroindoline-a and puroindoline-b) and non specific lipid transfer protein (ns-LTP1e1)ofTriticum aestivum seeds. Relationships with their in vitro antifungal properties. Plant Sci 138: 121–135.CrossRefGoogle Scholar
  31. Dubreil L, Meliande S, Chiron H, Compoint J-P, Quillien L, Branlard G, Marion D, 1998b. Effect of puroindolines on the bread making properties of wheat flour. Cereal Chem 75: 222–229.CrossRefGoogle Scholar
  32. Evrard A, Meynard D, Guiderdoni E, Joudrier P, Gautier M-F, 2007. The promoter of the wheat puroindoline-a gene (PinA) exhibits a more complex pattern of activity than that of thePinB gene and is induced by wounding and pathogen attack in rice. Planta 255: 287–300.Google Scholar
  33. Faize M, Sourice S, Dupuis F, Parisi L, Gautier M-F, Chevreau E, 2004. Expression of wheat puroindoline-b reduces scab susceptibility in transgenic apple (Malus × domestica Borkh.). Plant Sci 167: 347–354.CrossRefGoogle Scholar
  34. Galanade AA, Tiwari R, Ammiraju JSS, Santra DK, Lagu MD, Rao VS, et al. 2001. Genetic analysis of kernel hardness in bread wheat using PCR-based markers. Theor Appl Genet 103: 601–606.CrossRefGoogle Scholar
  35. Gautier M-F, Aleman M-E, 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.CrossRefPubMedGoogle Scholar
  36. Gautier M-F, Cosson P, Guirao A, Alary R, Joudrier P, 2000. Puroindoline genes are highly conserved in diploid ancestor wheats and related species but absent in tetraploidTriticum species. Plant Sci 153: 81–91CrossRefGoogle Scholar
  37. 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.CrossRefPubMedGoogle Scholar
  38. Gazza L, Conti S, Taddei F, Pogna NE, 2006. Molecular characterization of puroindolines and their encoding genesin Aegilops ventricosa. Mol Breed 17: 191–200.CrossRefGoogle Scholar
  39. Gazza L, Taddei F, Corbellini M, Cacciatori P, Pogna NE, 2008. Genetic and environmental factors affecting grain texture in common wheat. J Cereal Sci 47: 52–58.CrossRefGoogle Scholar
  40. Gedye KR, Morris CF, Bettge AD, 2004. Determination and evaluation of the sequence and textural effects of the puroindoline a and b genes in a population of synthetic hexaploid wheat. Theor Appl Genet 109: 1597–1603.CrossRefPubMedGoogle Scholar
  41. 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: 857–864.CrossRefGoogle Scholar
  42. Giroux MJ, Morris CF, 1998. Wheat grain hardness results from highly conserved mutations in the friabilin components puroindoline a and b. Proc Natl Acad Sci USA 95: 6262–6266.CrossRefPubMedGoogle Scholar
  43. 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 40: 370–374.CrossRefGoogle Scholar
  44. Giroux MJ, Sripo T, Gerhardt S, Sherwood J, 2003. Puroindolines: Their role in grain hardness and plant defense. In: Harding SE, ed. Biotechnology and Genetic Engineering Reviews 20: 277–290. Intercept, Andover, Hampshire, UK.Google Scholar
  45. Greenblatt GA, Bettge AD, Morris CF, 1995. The relationship among endosperm texture, friabilin occurrence, and bound polar lipids on wheat starch. Cereal Chem 72: 172–176.Google Scholar
  46. Greenwell P, Schofield JD, 1986. A starch granule protein associated with endosperm softness in wheat. Cereal Chem 63: 379–380.Google Scholar
  47. Guo SH, He ZH, Xia LQ, Wang HG, Zhang QZ, 2004. Detection of allelic variation for grain hardness in Chinese spring wheat by STS marker. Sci Agric Sin 37: 1797–1803.Google Scholar
  48. Hogg AC, Sripo 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.CrossRefPubMedGoogle Scholar
  49. Igrejas G, Gaborit T, Oury F-X, Chiron H, Marion D, Branlard G, 2001. Genetic and environmental effects on puroindoline-a and puroindoline-b content and their relationship to technological properties in French bread wheats. J Cereal Sci 34: 37–47.CrossRefGoogle Scholar
  50. Igrejas G, Leroy P, Charmet G, Gaborit T, Marion D, Branlard G, 2002. Mapping QTLs for grain hardness and puroindoline content in wheat (Triticum aestivum L.). Theor Appl Genet 106: 19–27.PubMedGoogle Scholar
  51. 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.CrossRefGoogle Scholar
  52. Jing W, Demcoe AR, Vogel AJ, 2003. Conformation of a bactericidal domain of puroindoline a: structure and mechanism of action of a 13-residue antimicrobial peptide. J Bacteriol 185: 4938–4947.CrossRefPubMedGoogle Scholar
  53. Jolly CJ, Rahman S, Kortt 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.CrossRefGoogle Scholar
  54. Krishnamurthy K, Balconi C, Sherwood JE, Giroux MJ, 2001a. Wheat puroindolines enhance fungal disease resistance in transgenic rice. Mol Plant Microbe In 14: 1255–1260.CrossRefGoogle Scholar
  55. Krishnamurthy K, Giroux MJ, 2001b. Expression of wheat puroindoline genes in transgenic rice enhances grain softness. Nat Biotechnol 19: 162–166.CrossRefPubMedGoogle Scholar
  56. Lawrence RJ, Pikaard CS, 2003. Transgene-induced RNA interference: a strategy for overcoming gene redundancy in polyploids to generate loss-of-function mutations. Plant J 36: 114–121.CrossRefPubMedGoogle Scholar
  57. Li G, He Z, Pena RJ, Xia X, Lillemo M, Qixin S, 2006. Identification of novel secaloindoline-a and secaloindoline-b alleles in CIMMYT hexaploid triticale lines. J Cereal Sci 43: 378–386.CrossRefGoogle Scholar
  58. Li W, Li H, Gill BS, 2008. Recurrent deletions of puroindoline genes At the grain Hardness locus in four independent lineages of polyploid wheat. Plant Physiol 146: 200–212.CrossRefPubMedGoogle Scholar
  59. 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.CrossRefGoogle Scholar
  60. Lillemo M, Ringlund K, 2002. Impact of puroindoline b alleles on the genetic variation for hardness in soft × hard wheat crosses. Plant Breed 121: 210–217.CrossRefGoogle Scholar
  61. Lillemo M, Simeone MC, Morris CF, 2002. Analysis of puroindoline a and b sequences fromTriticum aestivum cv. “Penawawa” and related diploid taxa. Euphytica 126: 321–331.CrossRefGoogle Scholar
  62. Luo L, Zhang J, Yang G, Li Y, Kexiu L, He G, 2008. Expression of puroindoline a enhances leaf rust resistance in transgenic tetraploid wheat. Mol Biol Rep 35: 195–200.CrossRefPubMedGoogle Scholar
  63. Martin CR, Rousser R, Brabec DL, 1993. Development of a single-kernel wheat characterization system. Transactions of the ASAE 36: 1399–1404.Google Scholar
  64. 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.CrossRefGoogle Scholar
  65. Martin J, Meyer F, Smidansky E, Wanjugi H, Blechl A, Giroux MJ, 2006. Complementation of the pina (null) allele with the wild typePina sequence restores a soft phenotype in transgenic wheat. Theor Appl Genet 113: 1563–1570.CrossRefPubMedGoogle Scholar
  66. Massa AN, Morris CF, Gill BS, 2004. Sequence diversity of puroindoline-a, puroindoline-b, and the grain softness protein inAegilops tauschii Coss. Crop Sci 44: 1808–1816.CrossRefGoogle Scholar
  67. McFadden ES, Sears ER, 1946. The origin ofTriticum spelta and its free-threshing hexaploid relatives. J Hered 37: 81–89, 107–116.PubMedGoogle Scholar
  68. McIntosh RA, Devos KM, Dubcovsky J, Rogers WJ, Morris CF, Appels R, Anderson OD, 2005. Catalogue of gene symbols for wheat: 2005 supplement. Scholar
  69. McIntosh RA, Devos KM, Dubcovsky J, Rogers WJ, Morris CF, Appels R, Somers DJ, Anderson OA, 2007. Catalogue of gene symbols for wheat: 2007 supplement. Scholar
  70. Morris CF, Bhave M, 2008. Reconciliation of D-genome puroindoline allele designations with current DNA sequence data. J Cereal Sci 48: 277–287.CrossRefGoogle Scholar
  71. Morris CF, Greenblatt GA, Bettge AD, Malkawi HI, 1994. Isolation and characterization of multiple forms of friabilin. J. Cereal Sci 21: 167–174.CrossRefGoogle Scholar
  72. Morris CF, King GE, Allan RE, Simeone MC, 2001a. Identification and characterization of near-isogenic hard and soft hexaploid wheats. Crop Sci 41: 211–217.CrossRefGoogle Scholar
  73. Morris CF, Lillemo M, Simeone MC, Giroux MJ, Babb SL, Kidwell KK, 2001b. Prevalence of puroindoline grain hardness genotypes among North American spring and winter wheats. Crop Sci 41: 218–228.CrossRefGoogle Scholar
  74. Morris CF, 2002. Puroindolines: the molecular genetic basis of wheat grain hardness. Plant Mol Biol 48: 633–647.CrossRefPubMedGoogle Scholar
  75. Perretant MR, Cadalen T, Charmet G, Sourdille P, Nicolas P, Boeuf C, et al. 2000. QTL analysis of bread-making quality in wheat using a doubled haploid population. Theor Appl Genet 100: 1167–1175.CrossRefGoogle Scholar
  76. Pogna NE, Gazza L, Korona V, Zanier R, Niglio A, Mei E, et al. 2002. Puroindolines and kernel hardness in wheat species. In: Ng PKW and Wrigley CW, eds. Wheat Quality Elucidation. AACC, St. Paul, Minnesota, USA, pp. 155–169.Google Scholar
  77. Rakszegi M, Wilkinson MD, Tosi P, Lovergrove A, Kovacs G, Bedo Z, Shewry PR, 2008. Puroindoline genes and proteins in tetraploid and hexaploid species ofTriticum. J Cereal Sci, doi:10.1016/j.jcs.2008.09.006.Google Scholar
  78. 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.Google Scholar
  79. Shewry PR, Jenkins J, Beaudoin F, Mills ENC, 2004. The classification, functions and evolutionary relationship 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
  80. Simeone MC, Lafiandra D, 2005. Isolation and characterization of friabilin genes in rye. J Cereal Sci 41: 115–122.CrossRefGoogle Scholar
  81. Slaughter DC, Norris KH, Hruschka WR, 1992. Quality and classification of hard red wheat. Cereal Chem 69: 428–432.Google Scholar
  82. Sourdille P, Perretant MR, Charmet G, Leroy P, Gautier M-F, Jourdier P, et al. 1996. Linkage between RFLP markers and genes affecting kernel hardness in wheat. Theor Appl Genet 93: 580–586.CrossRefGoogle Scholar
  83. Swan CG, Meyer FD, Hgg AC, Martin JM, Giroux, MJ, 2006. Puroindoline b limits binding of puroindoline a to starch and grain softness. Crop Sci 46: 1656–1665.CrossRefGoogle Scholar
  84. Symes KJ, 1965. The inheritance of grain hardness in wheat as measured by particle size index. Aust J Agric Res 16: 113–123.CrossRefGoogle Scholar
  85. Tanchak MA, Schernthaner JP, Giband M, Altosaar I, 1998. Tryptophanins: isolation and molecular characterization of oat cDNA clones encoding proteins structurally related to puroindoline and wheat grain softness proteins. Plant Sci 137: 173–184.CrossRefGoogle Scholar
  86. 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.CrossRefGoogle Scholar
  87. Turnbull KM, Gaborit T, Marion D, Rahman S, 2000. Variation in puroindoline polypeptides in Australian wheat cultivars in relation to grain hardness. Aust J Plant Physiol 27: 153–158.Google Scholar
  88. Turnbull KM, Turner M, Mukai Y, Yamamoto M, Morell MK, Appels R, Rahman S, 2003. The organization of genes tightly linked to theHa locus inAegilops tauschii, the D-genome donor to wheat. Genome 46: 330–338.CrossRefPubMedGoogle Scholar
  89. Turner AS, Bradburne RP, Fish L, Snape JW, 2004. New quantitative trait loci influencing grain texture and protein content in bread wheat. J Cereal Sci 40: 51–60.CrossRefGoogle Scholar
  90. Weightman RM, Millar S, Alava J, Foulkes MJ, Fish L, Snape JW, 2008. Effects of drought and the presence of the 1BL/1RS translocation on grain vitreosity, hardness and protein content in winter wheat. J Cereal Sci 47: 457–468.CrossRefGoogle Scholar
  91. Wilde PJ, Clark DC, Marion D, 1993. Influence of competitive adsorption of a lysopalmitoyl-phosphatidylcholine on the functional properties of puroindoline, a lipid-binding protein isolated from wheat flour. J Agric Food Chem 41: 1570–1576.CrossRefGoogle Scholar
  92. Wiley PR, Tosi P, Evrard A, Lovergrove A, Jones HD, Shewry PR, 2007. Promoter analysis and immunolocalisation show that puroindoline genes are exclusively expressed in starchy endosperm cells of wheat grain. Plant Mol Biol 64: 125–136.CrossRefPubMedGoogle Scholar
  93. Wilkinson M, Wan Y, Tosi P, Leverington M, Snape J, Mitchell RAC, Shewry PR, 2008. Identyfication and genetic mapping of variant forms of puroindoline b expressed in developing wheat grain. J Cereal Sci 48: 722–728.CrossRefGoogle Scholar
  94. Xia L, Chen F, He Z, Chen X, Morris CF, 2005. Occurrence of puroindoline alleles in Chinese winter wheats. Cereal Chem 82: 38–43.CrossRefGoogle Scholar
  95. Xia L, Geng H, Chen X, He Z, Lillemo M, Morris CF, 2008. Silencing of puroindoline a alters the kernel texture in transgenic bread wheat. J Cereal Sci 47: 331–338.CrossRefGoogle Scholar
  96. Ziemann M, Ramalingam A, Bhave M, 2008. Evidence of physical interactions of puroindoline proteins using the yeast two-hybrid system. Plant Sci 175: 307–313.CrossRefGoogle Scholar

Copyright information

© Institute of Plant Genetics, Polish Academy of Sciences, Poznan 2009

Authors and Affiliations

  1. 1.Plant Transformation and Cell Engineering DepartmentPlant Breeding and Acclimatization InstituteBłoniePoland

Personalised recommendations