Genetics and Applied Genomics of Quality Protein Maize for Food and Nutritional Security

  • P. K. Agrawal
  • M. G. Mallikarjuna
  • H. S. Gupta


Maize (Zea mays L.) is an important food and feed crop of the world. Together with rice and wheat, it provides around 40% of the food calories to more than 4.5 billion people in 94 developing countries. It also provides nearly 50% of the dietary protein for humans. In Africa and some of the Asian countries, almost 90% of maize grown is for human consumption and may account for 80–90% of the energy intake. In India, it is the third most important food crop after rice and wheat, both in terms of area and production. India is the fifth largest producer of maize in the world contributing 3% of the total global production. Protein malnutrition is widespread in the developing and underdeveloped countries, where 780 million people are affected by the same. Maize is the leading cereal in terms of production and accounts for 15% of proteins and 20% of calories requirement of the world. Protein malnutrition is caused by lack of access to adequate quantity and better quality protein intake and usually affects children and elderly persons. Maize, however, lacks adequate amounts of the essential amino acids, namely, lysine and tryptophan. Decades of efforts by maize researchers lead to the development of nutritionally superior maize cultivar popularly called as quality protein maize (QPM), which has twice the amount of lysine and tryptophan, thus making its quality as good as casein of milk. The o2 allele along with modifiers for tryptophan and lysine content and grain hardness made QPM agronomically suitable for cultivations. Intensive efforts were made by many workers to understand the genetics, molecular mechanism of QPM modifiers and applied these genomics knowledge to developed MAS-based QPM inbreds and commercial hybrids. All those studies and concerted efforts led to development and utilization of QPM. The area under QPM globally is more than 9.0 million hectares. Several reports were available on positive impact of QPM on children and adults. It has also been demonstrated in poultry and piggery, resulting in increased egg production and egg quality parameters and body mass. The area under QPM and consumption of QPM can be increased significantly by providing policy supports for QPM.


Lysine Malnutrition MAS Maize Nutrition Policy Protein QPM 


  1. Abiose SH, Ikujenlola AV, Abioderin FI (2015) Nutritional quality assessment of complementary foods produced from fermented and malted quality protein maize fortified with soybean flour. Polish J Food Nutr Sci 65:49–56CrossRefGoogle Scholar
  2. Agrawal PK, Gupta HS (2010) Enhancement of protein quality of maize using biotechnological options. Anim Nutr Feed Technol 10:79–91Google Scholar
  3. Agrawal PK, Babu BK, Saini N (2015) Omics of model plants. In: Plant omics: the omics of plant science. Springer India, New Delhi, pp 1–32Google Scholar
  4. Akalu G, Taffesse S, Gunaratna N, De Groote H (2010) The effectiveness of quality protein maize in improving the nutritional status of young children in the Ethiopian highlands. Food Nutr Bull 31:418–430CrossRefPubMedGoogle Scholar
  5. Anon (2008) Nutritious maize boosts growth of children in rural Ethiopia. African Science News ServiceGoogle Scholar
  6. Babu R, Prasanna BM (2014) Molecular breeding for quality protein maize (QPM). In: Genomics of plant genetic resources. Springer Netherlands, Dordrecht, pp 489–505CrossRefGoogle Scholar
  7. Babu R, Nair SK, Kumar A et al (2005) Two-generation marker-aided backcrossing for rapid conversion of normal maize lines to quality protein maize (QPM). Theor Appl Genet 111:888–897. CrossRefPubMedGoogle Scholar
  8. Babu BK, Agrawal PK, Saha S, Gupta HS (2015) Mapping QTLs for opaque2 modifiers influencing the tryptophan content in quality protein maize using genomic and candidate gene-based SSRs of lysine and tryptophan metabolic pathway. Plant Cell Rep 34:37–45. CrossRefPubMedGoogle Scholar
  9. Bai X (2002) Nutritional evaluation and utilization of quality protein maize Zhong Dan 9409 in broilers feed. Chinese Academy of Agricultural Sciences, BeijingGoogle Scholar
  10. Bantte K, Prasanna BM (2003) Simple sequence repeat polymorphism in quality protein maize (QPM) lines. Euphytica 129:337–344. CrossRefGoogle Scholar
  11. Belousov AA (1987) Genetic analysis of modified endosperm texture in opaque-2 maize. Sov Genet 23:459–464Google Scholar
  12. Bjarnason M, Vasal SK (1992) Breeding of quality protein maize (QPM). In: Janick J (ed) Plant breeding reviews. Wiley, New York, pp 181–216Google Scholar
  13. Black RE, Allen LH, Bhutta ZA et al (2008) Maternal and child undernutrition: global and regional exposures and health consequences. Lancet 371:243–260. CrossRefPubMedGoogle Scholar
  14. Bressani R (1990) Chemistry, technology, and nutritive value of maize tortillas. Food Rev Int 6:225–264. CrossRefGoogle Scholar
  15. Bressani R (1992) Nutritional value of high-lysine maize in humans. In: Mertz ET (ed) Quality protein maize. American Association of Cereal Chemists, St. Paul, pp 205–225Google Scholar
  16. Burnett RJ, Larkins BA (1999) Opaque2 modifiers alter transcription of the 27-kDa γ-zein genes in maize. Mol Gen Genet 261:908–916. CrossRefPubMedGoogle Scholar
  17. Burr FA, Burr B (1982) Three mutations in Zea mays affecting zein accumulation: a comparison of zein polypeptides, in vitro synthesis and processing, mRNA levels, and genomic organization. J Cell Biol 94:201–206. CrossRefPubMedGoogle Scholar
  18. Chopra N, Bhargawa A, Kumar A (2011) Effect of feeding quality protein maize (QPM) on growth of young children (1–3 years). Food Sci Res J 2:173–178Google Scholar
  19. Coleman CE, Larkins BA (1999) The prolamins of maize. In: Shewry PR, Casey R (eds) Seed proteins. Springer Netherlands, Dordrecht, pp 109–139CrossRefGoogle Scholar
  20. Coleman CE, Lopes MA, Gillikin JW et al (1995) A defective signal peptide in the maize high-lysine mutant floury 2. Proc Natl Acad Sci U S A 92:6828–6831. CrossRefPubMedPubMedCentralGoogle Scholar
  21. Coleman CE, Clore AM, Ranch JP et al (1997) Expression of a mutant a-zein creates the floury2 phenotype in transgenic maize. Proc Natl Acad Sci U S A 94:7094–7097. CrossRefPubMedPubMedCentralGoogle Scholar
  22. Dado RG (1999) Nutritional benefits of specially maize grain hybrids in dairy diets. J Anim Sci 77(Suppl):197–207CrossRefPubMedGoogle Scholar
  23. Damerval C, De Vienne D (1993) Quantification of dominance for proteins pleiotropically affected by opaque-2 in maize. Heredity (Edinb) 70:38–51. CrossRefGoogle Scholar
  24. Dannenhoffer JM, Bostwick DE, Or E, Larkins BA (1995) Opaque-15, a maize mutation with properties of a defective opaque-2 modifier. Proc Natl Acad Sci 92:1931–1935. CrossRefPubMedGoogle Scholar
  25. De Groote H, Kimenju SC, Morawetz UB (2011) Estimating consumer willingness to pay for food quality with experimental auctions: the case of yellow versus fortified maize meal in Kenya. Agric Econ 42:1–16. CrossRefGoogle Scholar
  26. De Groote H, Gunaratna NS, Fisher M et al (2016) The effectiveness of extension strategies for increasing the adoption of biofortified crops: the case of quality protein maize in East Africa. Food Secur 8:1101–1121. CrossRefGoogle Scholar
  27. De Paula H, Santos RC, Silva ME et al (2004) Biological evaluation of a nutritional supplement prepared with QPM maize cultivar BR 473 and other traditional food items. Brazilian Arch Biol Technol 47:247–251CrossRefGoogle Scholar
  28. De Steur H, Blancquaert D, Strobbe S et al (2015) Status and market potential of transgenic biofortified crops. Nat Biotechnol 33:25–29. CrossRefPubMedGoogle Scholar
  29. De-quan S, Shihuang Z (1994) Maize production and QPM breeding program in China. In: Larkins BA, Mertz ET (eds) Quality protein maize: 1964–1994. Proceedings of the international symposium on quality protein maize. 1–3 Dec EMBRAPA/CNPMS, Sete Lagoas, pp 108–123Google Scholar
  30. Dhillon BS, Prasanna BM (2001) Maize. In: Chopra VL (ed) Breeding field crops. Oxford and IBH, New Delhi, pp 147–189Google Scholar
  31. Di Fonzo N, Gentinetta E, Salamini F, Soave C (1979) Action of the opaque-7 mutation on the accumulation of storage products in maize endosperm. Plant Sci Lett 14:345–354. CrossRefGoogle Scholar
  32. Dreher K, Khairallah M, Ribaut J, Morris M (2003) Money matters (I): costs of field and laboratory procedures associated with conventional and marker-assisted maize breeding at CIMMYT. Mol Breed 11:221–234. CrossRefGoogle Scholar
  33. Emerson RA, Beadle GW, Fraser AC (1935) A summary of linkage studies in maize. Cornell Univ Agric Exp Stn Mem 180:1–83Google Scholar
  34. FAO (2015) (Accessed on Dec 2017)
  35. Gao J (2002) Nutritional evaluation and utilization of quality protein maize Zhong Dan 9409 in pig feed. Chinese Academy of Agricultural Sciences, BeijingGoogle Scholar
  36. Geetha KB, Lending CR, Lopes MA, et al. (1991) opaque-2 modifiers increase gamma-zein synthesis and alter its spatial distribution in maize endosperm. Plant Cell, 3:1207–1219.
  37. Geevers HO, Lake JK (1992) Development of modified opaque2 maize in South Africa. In: Mertz ET (ed) Quality protein maize. American Association of Cereal Chemists, St. Paul, pp 49–78Google Scholar
  38. Gibbon BC, Larkins BA (2005) Molecular genetic approaches to developing quality protein maize. Trends Genet 21:227–233. CrossRefPubMedGoogle Scholar
  39. Gibbon BC, Wang X, Larkins BA (2003) Altered starch structure is associated with endosperm modification in quality protein maize. Proc Natl Acad Sci 100:15329–15334. CrossRefPubMedGoogle Scholar
  40. Gillikin JW, Zhang F, Coleman CE et al (1997) A defective signal peptide tethers the floury-2 zein to the endoplasmic reticulum membrane. Plant Physiol 114:345–352. CrossRefPubMedPubMedCentralGoogle Scholar
  41. Gunaratna NS, De Groote H, Nestel P et al (2010) A meta-analysis of community-based studies on quality protein maize. Food Policy 35:202–210. CrossRefGoogle Scholar
  42. Gupta HS, Agrawal PK, Mahajan V et al (2009) Quality protein maize for nutritional security: rapid development of short duration hybrids through molecular marker assisted breeding. Curr Sci 96:230–237Google Scholar
  43. Gupta HS, Raman B, Agrawal PK et al (2013) Accelerated development of quality protein maize hybrid through marker-assisted introgression of opaque-2 allele. Plant Breed 132:77–82. CrossRefGoogle Scholar
  44. Gupta HS, Hossain F, Muthusamy V (2015a) Biofortification of maize: an Indian perspective. Indian J Genet Plant Breed 75:1–22. CrossRefGoogle Scholar
  45. Gupta HS, Hossain F, Nepolean T et al (2015b) Understanding genetic and molecular bases of Fe and Zn accumulation towards development of micronutrient-enriched maize. In: Rakshit A et al (eds) Nutrient use efficiency: from basics to advances. Springer India, New Delhi. CrossRefGoogle Scholar
  46. Habben JE, Kirleis AW, Larkins BA (1993) The origin of lysine-containing proteins in opaque-2 maize endosperm. Plant Mol Biol 23:825–838. CrossRefPubMedGoogle Scholar
  47. Hartings H, Lauria M, Lazzaroni N et al (2011) The Zea mays mutants opaque-2 and opaque-7 disclose extensive changes in endosperm metabolism as revealed by protein, amino acid, and transcriptome-wide analyses. BMC Genomics.
  48. Holding DR, Hunter BG, Chung T et al (2008) Genetic analysis of opaque2 modifier loci in quality protein maize. Theor Appl Genet 117:157–170. CrossRefPubMedGoogle Scholar
  49. Holding DR, Meeley RB, Hazebroek J et al (2010) Identification and characterization of the maize arogenate dehydrogenase gene family. J Exp Bot 61:3663–3673. CrossRefPubMedPubMedCentralGoogle Scholar
  50. Holding DR, Hunter BG, Klingler JP et al (2011) Characterization of opaque2 modifier QTLs and candidate genes in recombinant inbred lines derived from the K0326Y quality protein maize inbred. Theor Appl Genet 122:783–794. CrossRefPubMedGoogle Scholar
  51. Hunter BG, Beatty MK, Singletary GW et al (2002) Maize opaque endosperm mutations create extensive changes in patterns of gene expression. Plant Cell 14:2591–2612. CrossRefPubMedPubMedCentralGoogle Scholar
  52. Ignjatovic-Micic D, Stankovic G, Markovic K et al (2008) Quality protein maize: QPM. Genetika 40:205–214. CrossRefGoogle Scholar
  53. Institute of Medicine (2006) Protein and amino acids. Dietary Reference Intakes: the essential guide to nutrient requirements. Washington, DC: National Academies Press. pp. 145–155.Google Scholar
  54. Kim CS, Gibbon BC, Gillikin JW et al (2006) The maize mucronate mutation is a deletion in the 16-kDa gamma-zein gene that induces the unfolded protein response. Plant J 48:440–451. CrossRefPubMedGoogle Scholar
  55. Krivanek AF, De Groote H, Gunaratna NS et al (2007) Breeding and disseminating quality protein maize (QPM) for Africa. African J Biotechnol 6:312–324. CrossRefGoogle Scholar
  56. Lee KH, Jones RA, Dalby A, Tsai CY (1976) Genetic regulation of storage protein content in maize endosperm. Biochem Genet 14:641–650. CrossRefPubMedGoogle Scholar
  57. Lin KR, Bockholt AJ, Smith JD (1997) Utilization of molecular probes to facilitate development of quality protein maize. Maize Genet Coop Newsl 71:22–23Google Scholar
  58. Liu H, Shi J, Sun C et al (2016) Gene duplication confers enhanced expression of 27-kDa γ-zein for endosperm modification in quality protein maize. Proc Natl Acad Sci 113:4964–4969. CrossRefPubMedGoogle Scholar
  59. Lopes MA, Larkins BA (1991) Gamma-zein content is related to endosperm modification in quality protein maize. Crop Sci 31:1655–1662CrossRefGoogle Scholar
  60. Ma Y, Nelson OE (1975) Amino acid composition and storage proteins in two new high-lysine mutants in maize. Cereal Chem 52:412–419Google Scholar
  61. Mallikarjuna MG (2015) Studies on genetics and genomics of kernel iron and zinc in maize (Zea mays L.). ICAR-Indian Agricultural Research Institute, New DelhiGoogle Scholar
  62. Mallikarjuna MG, Nepolean T, Hossain F et al (2014) Genetic variability and correlation of kernel micronutrients among exotic quality protein maize inbreds and their utility in breeding programme. Indian J Genet Plant Breed.
  63. Mallikarjuna MG, Thirunavukkarasu N, Hossain F et al (2015) Stability performance of inductively coupled plasma mass spectrometry-phenotyped kernel minerals concentration and grain yield in maize in different agro-climatic zones. PLoS One.
  64. Mamatha H, Meena MK, Kumar PC (2017) Quality protein maize (QPM) as balance nutrition for human diet. Adv Plants Agric Res 6:5–6. CrossRefGoogle Scholar
  65. Maseta EJ (2016) Efficacy of quality protein maize-based supplementary foods on rehabilitating undernourished children in Mvomero District, Tanzania. Sokoine University of Agriculture, MorogoroGoogle Scholar
  66. Mertz ET, Bates LS, Nelson OE (1964) Mutant gene that changes protein composition and increases lysine content of maize endosperm. Science 145:279–280. CrossRefPubMedGoogle Scholar
  67. Misra PS, Jambunathan R, Mertz ET et al (1972) Endosperm protein synthesis in maize mutants with increased lysine content. Science 176:1425–1427. CrossRefPubMedGoogle Scholar
  68. Mpofu ID, Sibanda S, Shonihwa A, Pixely K (2012) The nutritional value of quality protein maize for weaner pigs. J Pet Environ Biotechnol 3:3–6. CrossRefGoogle Scholar
  69. Myers AM, James MG, Lin Q et al (2011) Maize opaque5 encodes monogalactosyldiacylglycerol synthase and specifically affects galactolipids necessary for amyloplast and chloroplast function. Plant Cell 23:2331–2347. CrossRefPubMedPubMedCentralGoogle Scholar
  70. Nedi G, Agriculture C, Medicine V, Box PO (2016) Review on quality protein maize breeding for ethiopia. J Biol Agric Healthc 6:84–96Google Scholar
  71. Nelson OE (1969) Genetic modification of protein quality in plants. Adv Agron 21:171–194CrossRefGoogle Scholar
  72. Nelson OE (1981) The mutants opaque-9 through opaque-13. Corn Genet Coop Newsl 55:68Google Scholar
  73. Nelson OE, Mertz ET, Bates LS (1965) Second mutant gene affecting the amino acid pattern of maize endosperm proteins. Science 150:1469–1470. CrossRefPubMedGoogle Scholar
  74. Nuss ET, Tanumihardjo SA (2011) Quality protein maize for Africa: closing the protein inadequacy gap in vulnerable populations. Adv Nutr An Int Rev J 2:217–224. CrossRefGoogle Scholar
  75. Omage JJ, Agubosi OCP, Bawa GS, Onimisi P (2009) Evaluation of the nutritive value of quality protein maize on the growth performance and carcass characteristics of weaner rabbits. Pakistan J Nutr 8:106–111CrossRefGoogle Scholar
  76. Ortega EI, Bates LS (1983) Biochemical and agronomic studies of two modified hard-endosperm opaque-2 maize (Zea mays L.) populations. Cereal Chem 60:107–111Google Scholar
  77. Osborne TB, Mendel LB (1914) Nutritive properties of proteins of the maize kernel. J Biol Chem 18:1–16Google Scholar
  78. Osei SA, Atuahene C, Okai DB et al (1998) The nutritive value of quality protein maize in the diets of broiler chickens. Ghana J Agric Sci 31:1–5Google Scholar
  79. Osei SA, Dei HK, Tuah AK (1999) Evaluation of quality protein maize as a feed ingredient for layer pullet. J Anim Feed Sci 8:181–189. CrossRefGoogle Scholar
  80. Panda AK, Raju MVLN, Rama Rao SV et al (2010) Replacement of normal maize with quality protein maize on performance, immune response and carcass characteristics of broiler chickens. Asian-Australasian J Anim Sci 23:1626–1631. CrossRefGoogle Scholar
  81. Panda AK, Raju MVLN, Rao SVR et al (2011) Nutritional evaluation and utilisation of quality protein maize, Nityashree hybrid maize, and normal maize in broiler chickens. Br Poult Sci 52:632–638. CrossRefPubMedGoogle Scholar
  82. Pfunde CN, Mutengwa CS (2016) Combining ability of quality protein maize inbred lines for seedling tolerance to drought stress. Philipp J Crop Sci 41:1–12Google Scholar
  83. Prandini A, Sigolo S, Morlacchini M et al (2011) High-protein maize in diets for growing pigs. Anim Feed Sci Technol 165:105–110. CrossRefGoogle Scholar
  84. Prasanna BM, Sarkar KR (1991) Coordinate genetic regulation of maize endosperm. Maize genetics perspectives, ICAR, pp 74–86Google Scholar
  85. Prasanna BM, Vasal SK, Kassahun B, Singh NN (2001) Quality protein maize. Curr Sci 81:1308–1319Google Scholar
  86. Prasanna BM, Pixley K, Warburton ML, Xie C-X (2010) Molecular marker-assisted breeding options for maize improvement in Asia. Mol Breed 26:339–356. CrossRefGoogle Scholar
  87. Ribaut J-M, Hoisington D (1998) Marker-assisted selection: new tools and strategies. Trends Plant Sci 3:236–239. CrossRefGoogle Scholar
  88. Rugema H (2014) Promotion of quality protein maize as a strategic solution to addressing food and nutrition security: the legacy of Dr. Wayne Haag. African J Food Agric Nutr Dev 14:1–9Google Scholar
  89. Salamini F, Fonzo NDI, Gentinetta E, Soave C (1979) A dominant mutation interfering with protein accumulation in maize seeds. In: Seed protein improvement in cereals and grain legumes. IAEA, Vienna, pp 97–108Google Scholar
  90. Salamini F, Di Fonzo N, Fornasari E et al (1983) Mucronate, Mc, a dominant gene of maize which interacts with opaque-2 to suppress zein synthesis. Theor Appl Genet 65:123–128. CrossRefPubMedGoogle Scholar
  91. Schmidt RJ, Burr FA, Burr B (1987) Transposon tagging and molecular analysis of the maize regulatory locus opaque-2. Science 238:960–963. CrossRefPubMedGoogle Scholar
  92. Schmidt RJ, Burrt FA, Aukerman MJ, Burr B (1990) Maize regulatory gene opaque-2 encodes a protein with a ‘leucine-zipper’ motif that binds to zein DNA. Proc Natl Acad Sci U S A 87:46–50. CrossRefPubMedPubMedCentralGoogle Scholar
  93. Shewry PR (2007) Improving the protein content and composition of cereal grain. J Cereal Sci 46:239–250. CrossRefGoogle Scholar
  94. Shiferaw B, Prasanna BM, Hellin J, Bänziger M (2011) Crops that feed the world 6. Past successes and future challenges to the role played by maize in global food security. Food Secur 3:307–327. CrossRefGoogle Scholar
  95. Singleton WR (1939) Recent linkage studies in maize: V. opaque endosperm-2 (2). Genetics 24:59–63Google Scholar
  96. Soave C, Tardani L, Di Fonzo N, Salamini F (1981) Zein level in maize endosperm depends on a protein under control of the opaque-2 and opaque-6 loci. Cell 27:403–410. CrossRefPubMedGoogle Scholar
  97. Sofi PA, Wani SA, Rather AG, Wani SH (2009) Quality protein maize (QPM): genetic manipulation for the nutritional fortification of maize. J Plant Breed Crop Sci 1:244–253Google Scholar
  98. Stevens R, Winter-Nelson A (2008) Consumer acceptance of provitamin A-biofortified maize in Maputo, Mozambique. Food Policy 33:341–351. CrossRefGoogle Scholar
  99. Tamir B, Gebrehawariat E, Tegegne A, Kortu MY (2012) Rumen degradability characteristics of normal maize stover and silage, and quality protein maize silage-based diets offered to cows. Trop Anim Health Prod 44:1547–1553. CrossRefPubMedGoogle Scholar
  100. Tandzi LN, Mutengwa CS, Ngonkeu ELM et al (2017) Breeding for quality protein maize (QPM) varieties: a review. Agronomy 7:80. CrossRefGoogle Scholar
  101. Thompson GA, Larkins BA (1994) Characterization of zein genes and their regulation in maize endosperm. In: Freeling M, Walbot V (eds) The maize handbook. Springer New York, New York, pp 639–647CrossRefGoogle Scholar
  102. Tsai CY, Dalby A (1974) Comparison of the effect of shrunken-4, opague-2, opaque-7, and floury-2 genes on the zein content of maize during endosperm development. Cereal Chem 51:825–828Google Scholar
  103. Vasal SK (2000) The quality protein maize story. Food Nutr Bull 21:445–450. CrossRefGoogle Scholar
  104. Vasal SK (2001) High quality protein corn. In: Hallauer AR (ed) Specialty corns. CRC Press, Boca Raton, Florida, USA, pp 93–137Google Scholar
  105. Vasal SK (2002) Quality protein maize: overcoming the hurdles. J Crop Prod 6:193–227. CrossRefGoogle Scholar
  106. Vasal SK, Villegas E, Bjarnason M et al (1980) Genetics modifiers and breeding strategies in developing hard endosperm. In: Pollmer WG, Phipps RH (eds) Improvement of quality traits of maize for grains and silage use. Nighoff, The Hague, pp 37–73Google Scholar
  107. Vaswani S, Kumar R, Kumar V (2015) In vitro nutritional evaluation of normal and quality protein maize fodders for ruminants. Indian J Anim Nutr 32:20–24Google Scholar
  108. Villegas E, Vasal SK, Bjarnason M (1992) Quality protein maize – what is it and how was it developed. In: Mertz ET (ed) Quality protein maize. American Association of Cereal Chemists, St. Paul, pp 27–48Google Scholar
  109. Vivek BS, Krivanek AF, Palacios-rojas N et al (2008) Breeding quality protein maize: protocols for developing QPM cultivars. CIMMYT, MexicoGoogle Scholar
  110. Wallace JC, Lopes MA, Paiva E, Larkins BA (1990) New methods for extraction and quantitation of zeins reveal a high content of gamma-zein in modified opaque-2 maize. Plant Physiol 92:191–196. CrossRefPubMedPubMedCentralGoogle Scholar
  111. Wang G, Sun X, Wang G et al (2011) Opaque7 encodes an acyl-activating enzyme-like protein that affects storage protein synthesis in maize endosperm. Genetics 189:1281–1295. CrossRefPubMedPubMedCentralGoogle Scholar
  112. Wang G, Qi W, Wu Q et al (2014) Identification and characterization of maize floury4 as a novel semidominant opaque mutant that disrupts protein body assembly. Plant Physiol 165:582–594. CrossRefPubMedPubMedCentralGoogle Scholar
  113. WHO (2007) Protein and amino acid requirements in human nutrition: report of a joint FAO/WHO/UNU expert consultation. WHO Technical Report Series no. 935. Available from:
  114. Wu Y, Holding DR, Messing J (2010) γ-zeins are essential for endosperm modification in quality protein maize. Proc Natl Acad Sci 107:12810–12815. CrossRefPubMedGoogle Scholar
  115. Xu Y, Crouch JH (2008) Marker-assisted selection in plant breeding: from publications to practice. Crop Sci 48:391–407. CrossRefGoogle Scholar
  116. Yang W, Zheng Y, Zheng W, Feng R (2005) Molecular genetic mapping of a high-lysine mutant gene (opaque-16) and the double recessive effect with opaque-2 in maize. Mol Breed 15:257–269. CrossRefGoogle Scholar
  117. Yang W, Zheng Y, Wu J (2008) Heterofertilization of the opaque-2 endosperm in maize. Hereditas 145:225–230. CrossRefPubMedGoogle Scholar
  118. Yuan L, Dou Y, Kianian SF et al (2014) Deletion mutagenesis identifies a haploinsufficient role for γ-zein in opaque2 endosperm modification. Plant Physiol 164:119–130. CrossRefPubMedGoogle Scholar
  119. Zaidi PH, Vasal SK, Maniselvan P et al (2008) Stability in performance of quality protein maize under abiotic stress. Maydica 53:249–260Google Scholar
  120. Zhai S (2002) Nutritional evaluation and utilization of quality protein maize Zhong Danm 9409 in laying hen feed. Northwestern Agricultural and Forestry University of Science and Technology, ShaanxiGoogle Scholar
  121. Zhang WL, Yang WP, Chen ZW et al (2010) Molecular marker-assisted selection for o2 introgression lines with o16 gene in corn. Acta Agron Sin 36:1302–1309. CrossRefGoogle Scholar
  122. Zhang W, Yang W, Wang M et al (2013) Increasing lysine content of waxy maize through introgression of opaque-2 and opaque-16 genes using molecular assisted and biochemical development. PLoS One 8:4–13. CrossRefGoogle Scholar
  123. Zhang Z, Zheng X, Yang J et al (2016) Maize endosperm-specific transcription factors O2 and PBF network the regulation of protein and starch synthesis. Proc Natl Acad Sci 113:10842–10847. CrossRefPubMedGoogle Scholar

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© Springer International Publishing AG, part of Springer Nature 2018

Authors and Affiliations

  • P. K. Agrawal
    • 1
  • M. G. Mallikarjuna
    • 2
  • H. S. Gupta
    • 3
  1. 1.National Agricultural Science Fund, Indian Council of Agricultural ResearchNew DelhiIndia
  2. 2.Division of GeneticsICAR-Indian Agricultural Research InstituteNew DelhiIndia
  3. 3.ICAR-Indian Agricultural Research InstituteNew DelhiIndia

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