Biofortification in Pearl Millet: From Conception to Dissemination

  • Alphonse Vinoth
  • Ramalingam RavindhranEmail author
Part of the Concepts and Strategies in Plant Sciences book series (CSPS)


Biofortification is an economical and sustainable process of delivering essential micronutrients through staple crops. The biofortified crops developed by HarvestPlus through conventional breeding continue to reach the target populations of Asia and Africa in order to reduce the burden of iron, zinc and vitamin A deficiency. Pearl millet, a dryland crop of the arid and semi-arid tropics is a suitable crop for iron biofortification as it harbours sufficient genetic variability for grain iron (Fe) and zinc (Zn) in the existing germplasm. Zn is highly correlated with grain Fe and therefore enhanced as an associated trait during the breeding for high-iron pearl millet. ICTP 8203 Fe-10-2, an iron-biofortified pearl millet (Fe-PM) variety developed via intra-population improvement of iniadi germplasm, was commercially released for cultivation in Maharashtra, India, by 2014. Efficacy trials undertaken in women and children feeding on Fe-PM meals revealed an enhancement in their micronutrient status as well as their functional outcomes. Disbursement of Fe-PM through public–private seed markets worked out to be cost-effective. Farmers readily adopted Fe-PM for cultivation based on its superior agronomic performance rather than the preference for consumer attributes. On the other hand, consumers expressed their willingness to pay for Fe-PM over regular pearl millet because of its favourable sensory characteristics. Therefore, investment on high-Fe hybrids would bridge the gap between the farmers and consumers acceptance of biofortified millets. Iron biofortification is also limited by the presence of antinutrients like phytates and polyphenols as they hinder the Fe bioavailability. The development of biofortified crops with reduced antinutrients needs careful evaluation as they have a significant role in protection against diseases and seedling growth. This review paper deliberately describes the success of high-Fe pearl millet in India and the lessons to be learnt for expanding the biofortification efforts to other small millets.


Biofortification Iron deficiency HarvestPlus Pearl millet Conventional breeding ICRISAT Dhanshakti Seed distribution system 



Atomic absorption spectroscopy


Consultative Group on International Agricultural Research


Disability-adjusted life year


Estimated average requirement






Iron-biofortified pearl millet

G × E





Inductively coupled plasma optical emission spectroscopy


International Crops Research Institute for the Semi-Arid Tropics


Low-phytate mutants


National Sample Survey Office


Open-pollinated variety


Phytic acid


State Agricultural University


United Nations International Children’s Emergency Fund


X-ray fluorescence spectroscopy





The authors acknowledge with sincere thanks the financial support received from Loyola College–Times of India Major Research Project Scheme (Project approval code: 4LCTOI14PBB001) and University Grants Commission, New Delhi Research Award Scheme (F.30-1/2014/RA-2014-16-GE-TAM-5825 SA-II), for their research on millet biofortification.

Conflict of Interest Statement: The authors declare that the manuscript was written in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.


  1. Abdalla AA, El Tinay AH, Mohamed BE, Abdalla AH (1998) Effect of traditional processes on phytate and mineral content of pearl millet. Food Chem 63(1):79–84CrossRefGoogle Scholar
  2. Adams CL, Hambidge M, Raboy V, Dorsch JA, Sian L, Westcott JL, Krebs NF (2002) Zinc absorption from a low-phytic acid maize. Am J Clin Nutr 76(3):556–559CrossRefGoogle Scholar
  3. Andrews DJ, Kumar KA (1992) Pearl millet for food, feed and forage. Adv Agron 48:89–139CrossRefGoogle Scholar
  4. Banerji A, Birol E, Karandikar B, Rampal J (2016) Information, branding, certification, and consumer willingness to pay for high-iron pearl millet: evidence from experimental auctions in Maharashtra, India. Food Policy 62:133–141CrossRefGoogle Scholar
  5. Basavaraj G, Rao PP, Bhagavatula S, Ahmed W (2010) Availability and utilization of pearl millet in India. SAT eJournal 8:1–6Google Scholar
  6. Bashir EMA, Ali AM, Ali AM, Melchinger AE, Parzies HK, Haussmann BIG (2014) Characterization of Sudanese pearl millet germplasm for agro-morphological traits and grain nutritional value. Plant Genet Res 12(1):35–47CrossRefGoogle Scholar
  7. Birol E, Asare-Marfo D, Karandikar B, Roy D (2011) A latent class approach to investigating farmer demand for biofortified staple food crops in developing countries: the case of high-iron pearl millet in Maharashtra, India. HarvestPlus working paper No.7, October 2011, pp 1–17Google Scholar
  8. Birol E, Meenakshi JV, Oparinde A, Perez S, Tomlins K (2015) Developing country consumer’s acceptance of biofortified foods: a synthesis. Food Secur 7(3):555–568CrossRefGoogle Scholar
  9. Boccio JR, Iyengar V (2003) Iron deficiency: causes, consequences, and strategies to overcome this nutritional problem. Biol Trace Elem Res 94(1):1–32CrossRefGoogle Scholar
  10. Bouis HE (1999) Economics of enhanced micronutrient density in food staples. Field Crop Res 60(1/2):165–173CrossRefGoogle Scholar
  11. Bouis HE (2000) Enrichment of food staples through plant breeding: a new strategy for fighting micronutrient malnutrition. Nutrition 16(7/8):701–704CrossRefGoogle Scholar
  12. Bouis HE (2003) Micronutrient fortification of plants through plant breeding: can it improve nutrition in man at low cost? Proc Nutr Sci 62(2):403–411CrossRefGoogle Scholar
  13. Bouis HE, Hotz C, McClafferty B, Meenakshi JV, Pfeiffer WH (2011) Biofortification: a new tool to reduce micronutrient malnutrition. Food Nutr Bull 32:31S–40SCrossRefGoogle Scholar
  14. Bregitzer P, Raboy V (2006) Effects of four independent low-phytate mutations on barley agronomic performance. Crop Sci 46(3):1318–1322CrossRefGoogle Scholar
  15. Burger A, Høgh-Jensen H, Gondah J, Hash CT, Haussmann BIG (2014) Micronutrient density and stability in West African pearl millet—potential for biofortification. Crop Sci 54(4):1709–1720CrossRefGoogle Scholar
  16. Caballero B (2002) Global patterns of child health: the role of nutrition. Ann Nutr Metab 46(1):3–7CrossRefGoogle Scholar
  17. Carvalho S, Vasconcelos MW (2013) Producing more with less: strategies and novel technologies for plant-based food biofortification. Food Res Int 54(1):961–971CrossRefGoogle Scholar
  18. Cercamondi CI, Egli IM, Mitchikpe E, Tossou F, Zeder C, Hounhouigan JD, Hurrell RF (2013) Total iron absorption by young women from iron-biofortified pearl millet composite meals is double that from regular millet meals but less than that from post-harvest iron-fortified millet meals. J Nutr 143:1376–1382CrossRefPubMedPubMedCentralGoogle Scholar
  19. Cherian B (2014) Delivery of pearl millet in India. Biofortification progress briefs, August 2014, pp 57–58Google Scholar
  20. Cichy KA, Forster S, Grafton KF, Hosfield GL (2005) Inheritance of seed zinc accumulation in navy bean. Crop Sci 45(3):864–870CrossRefGoogle Scholar
  21. Dahlberg JA, Wilson JP, Snyder T (2004) Sorghum and pearl millet: health foods and industrial products in developed countries. In: Alternative uses of sorghum and pearl millet in Asia. Proceedings of the expert meeting, ICRISAT, Patancheru, Andhra Pradesh, India, 1–4 July 2003.CFC tech paper 34, pp 42–59Google Scholar
  22. De Moura FF, Palmer A, Finkelstein J, Haas JD, Murray-Kolb LE, Wenger MJ, Birol E, Boy E, Peña-Rosas JP (2014) Are biofortified staple food crops improving vitamin A and iron status in women and children? New evidence from efficacy trials. Adv Nutr 5(5):568–570CrossRefPubMedPubMedCentralGoogle Scholar
  23. FAO (2003) The state of food insecurity in the world. Food and Agricultural Organization of the United Nations, Rome/GenevaGoogle Scholar
  24. Finkelstein JL, Mehta S, Udipi SA, Ghugre PS, Luna SV, Wenger MJ, Murray-Kolb LE, Przybyszewski EM, Haas JD (2015) A randomized trial of iron-biofortified pearl millet in school children in India. J Nutr 145(7):1576–1581CrossRefGoogle Scholar
  25. Gangashetty PI, Motagi BN, Pavan R, Roodagi MB (2016) Breeding crop plants for improved human nutrition through biofortification: progress and prospects. In: Al-Khayri JM, Jain SM, Johnson DV (eds) Advances in plant breeding strategies: agronomic, abiotic and biotic stress traits. Springer, Switzerland, pp 35–76CrossRefGoogle Scholar
  26. Gibson RS, Donavan UM, Heath MAL (1994) Dietary strategies to improve the iron and zinc nutriture of young women following a vegetarian diet. Plant Foods Hum Nutr 51(1):1–16CrossRefGoogle Scholar
  27. Gomez-Becerra HF, Erdem H, Yazici A, Tutus Y, Torunb B, Ozturk L (2010) Grain concentrations of protein and mineral nutrients in a large collection of spelt wheat grown under different environments. J Cereal Sci 52(3):342–349CrossRefGoogle Scholar
  28. Govindaraj M, Selvi B, Rajarathinam S (2009) Correlation studies for grain yield components and nutritional quality traits in pearl millet (Pennisetum glaucum (L.) R. Br.) germplasm. World J Agric Sci 5(6):686–689Google Scholar
  29. Govindaraj M, Selvi M, Rajarathinam S, Sumathi P (2011) Genetic variability, heritability and genetic advance in India’s pearl millet (Pennisetum glaucum (L) R. Br.) accessions for yield and nutritional quality traits. Afr J Food Agric Nutr Dev 11(3):4758–4771Google Scholar
  30. Govindaraj M, Rai KN, Shanmugasundaram P (2013) Combining ability and heterosis for grain iron and zinc density in pearl millet. Crop Sci 53(2):507–517CrossRefGoogle Scholar
  31. Graham RD, Welch RM, Bouis HE (2001) Addressing micronutrient malnutrition through enhancing the nutritional quality of staple foods: principles, perspectives and knowledge gap. Adv Agron 70:77–142CrossRefGoogle Scholar
  32. Gupta SK, Velu G, Rai KN, Sumalini K (2009) Association of grain iron and zinc content with grain yield and other traits in pearl millet (Pennisetum glaucum (L.) R. BR). Crop Improv 36(2):4–7Google Scholar
  33. Gupta SK, Rai KN, Singh P, Ameta VL, Gupta SK, Jeyalekha AK, Mahala RS, Pareek S, Swami ML, Verma YS (2015) Seed set variability under high temperatures during flowering period in pearl millet (Pennisetum glaucum L. (R.) Br.). Field Crops Res 171:41–53CrossRefGoogle Scholar
  34. Haas JE (2014) Efficacy and other nutrition evidence for iron crops. Biofortification progress briefs, August 2014, pp 39–40Google Scholar
  35. Haas JD, Brownlie T (2001) Iron deficiency and reduced work capacity: a critical review of the research to determine a causal relationship. J Nutr 131(Suppl):676S–88SCrossRefGoogle Scholar
  36. Hambidge MK, Miller LV, Westcott JE, Sheng X, Krebs NF (2010) Zinc bioavailability and homeostasis. Am J Clin Nutr 91(5):1478S–1483SCrossRefPubMedPubMedCentralGoogle Scholar
  37. Hirschi K (2009) Nutrient biofortification of food crops. Ann Rev Nutr 29:401–421CrossRefGoogle Scholar
  38. Hotz C (2013) Biofortification. In: Caballero B, Allen L, Prentice A (eds) Encyclopedia of human nutrition, vol 1. Elsevier, Oxford, UK, pp 175–181CrossRefGoogle Scholar
  39. Hotz C, McClafferty B (2007) From harvest to health: challenges for developing biofortified staple foods and determining their impact on micronutrient status. Food Nutr Bull 28(2 suppl):S271–S279CrossRefGoogle Scholar
  40. Huey SL, Venkatramanan S, Udipi SA, Finkelstein JL, Ghugre P, Haas JD, Thakker V, Thorat A, Salvi A, Kurpad AV, Mehta S (2017) Acceptability of iron- and zinc-biofortified pearl millet (ICTP-8203)-based complementary foods among children in an urban slum of Mumbai, India. Front Nutr 4:39CrossRefPubMedPubMedCentralGoogle Scholar
  41. Hurrell RF (2003) Influence of vegetable protein sources on trace element and mineral bioavailability. J Nutr 133(9):S2973–S2977CrossRefGoogle Scholar
  42. ICMR (2009) Nutrient requirements and recommended dietary allowances for Indians: a report of the expert group of the Indian Council of Medical Research 2009. National Institute of Nutrition, HyderabadGoogle Scholar
  43. Kanatti A, Rai K, Radhika K, Govindaraj M, Sahrawat KL, Rao AS (2014) Grain iron and zinc density in pearl millet: combining ability, heterosis and association with grain yield and grain size. SpringerPlus 3:763CrossRefPubMedPubMedCentralGoogle Scholar
  44. Kodkany BS, Bellad RM, Mahantshetti NS, Westcott JE, Krebs NF, Kemp JF, Hambidge KM (2013) Biofortification of pearl millet with iron and zinc in a randomized controlled trial increases absorption of these minerals above physiologic requirements in young children. J Nutr 143(9):1489–1493CrossRefPubMedPubMedCentralGoogle Scholar
  45. Krishnaswamy K (2009) The problem and consequences of the double burden—a brief overview. In: Programme and abstracts. Symposium on nutritional security for India—issues and way forward, Indian National Science Academy, New Delhi, pp 5–6Google Scholar
  46. Lemke S (2005) Nutrition security, livelihoods and HIV/AIDS: implications for research among farm worker households in South Africa. Public Health Nutr 8(7):844–852CrossRefGoogle Scholar
  47. Lestienne I, Icardnière C, Mouquet C, Picq C, Trèche S (2005) Effect of soaking whole cereal and legume seeds on iron, zinc and phytate contents. Food Chem 89(3):421–425CrossRefGoogle Scholar
  48. Liu ZH, Wang HY, Wang XE, Zhang GP, Chen PD, Liu DJ (2006) Genotypic and spike positional difference in grain phytase activity, phytate, inorganic phosphorus, iron, and zinc contents in wheat (Triticum aestivum L.). J Cereal Sci 44(2):212–219Google Scholar
  49. Lönnerdal B (2002) Phytic acid-trace element (Zn, Cu, Mn) interactions. Int J Food Sci Technol 37(7):749–758CrossRefGoogle Scholar
  50. Mayer JE, Pfeiffer WH, Beyer P (2008) Biofortified crops to alleviate micronutrient malnutrition. Plant Biol 11(2):166–170Google Scholar
  51. Meenakshi JV, Nancy J, Manyong V, De Groote H, Javelosa J, Yanggen D, Naher F, Garcia J, Gonzales C, Ming E (2010) How cost effective is biofortification in combating micronutrient malnutrition? An ex ante assessment. World Dev 38(1):64–75CrossRefGoogle Scholar
  52. Mendoza C (2002) Effect of genetically modified low phytic acid plants on mineral absorption. Int J Food Sci Technol 37(7):759–767CrossRefGoogle Scholar
  53. Nestel P, Bouis H, Meenakshi JV, Pfeiffer W (2006) Biofortification of staple food crops. J Nutr 136(4):1064–1067CrossRefGoogle Scholar
  54. Pfeiffer W, McClafferty B (2007) Biofortification: breeding micronutrient-dense crops. In: Kang MS, Priyadarshan PM (eds) Breeding major food staples. Blackwell Publishing, Ames, pp 61–91CrossRefGoogle Scholar
  55. Raboy V (2003) myo-Inositol-1–6-hexakisphosphate. Phytochemistry 64(6):1033–1043CrossRefGoogle Scholar
  56. Raboy V (2007) Seed phosphorus and the development of low-phytate crops. In: Turner BL, Richardson AE, Mullaney EJ (eds) Inositol phosphates: linking agriculture and the environment. CABI, Wallingford, UK, pp 111–132Google Scholar
  57. Rai KN, Anand Kumar K, Andrews DJ, Rao AS, Raj AGB, Witcombe JR (1990) Registration of ICTP 8203 pearl millet. Crop Sci 30:959CrossRefGoogle Scholar
  58. Rai KN, Yadav OP, Rajpurohit BS, Patil HT, Govindaraj M, Khairwal IS, Rao AS, Shivade H, Pawar VY, Kulkarni MP (2013) Breeding pearl millet cultivars for high iron density with zinc density as an associated trait. J SAT Agric Res 11:1–7Google Scholar
  59. Rai KN, Patil HT, Yadav OP, Govindaraj M, Khairwal IS, Cherian B, Rajpurohit BS, Rao AS, Shivade H, Kulkarni MP (2014) Dhanashakti: a high-iron pearl millet variety. Indian Farming 64(7):32–34Google Scholar
  60. Rai K (2014) Iron pearl millet. Biofortification progress briefs, August 2014, pp 7–8Google Scholar
  61. Reddy BVS, Ramesh S, Longvah T (2005) Prospects of breeding for micronutrients and β-carotene-dense sorghums. Int Sorghum Millets Newslett 46:10–14Google Scholar
  62. Rosegrant MW, Cline SA (2003) Global food security: challenges and policies. Science 302(5652):1917–1919CrossRefGoogle Scholar
  63. Saltzman A, Birol E, Bouis H, Boy E, De Moura FF, Islam Y, Pfeiffer WH (2013) Biofortification: progress toward a more nourishing future. Glob Food Secur 2(1):9–17CrossRefGoogle Scholar
  64. Sehgal S, Kawatra A, Singh G (2004) Recent advances in pearl millet and sorghum processing and food product development. In: Alternative uses of sorghum and pearl millet in Asia. Proceedings of the expert meeting, ICRISAT, Patancheru, Andhra Pradesh, India, 1–4 July 2003. CFC technical paper no. 34, pp 60–92Google Scholar
  65. Shafii B, Price WJ (1998) Analysis of genotype-by environment interaction using the additive main effects and multiplicative interaction model and stability estimates. J Agric Biol Environ Stat 3:335–345CrossRefGoogle Scholar
  66. Shivran AC (2016) Biofortification for nutrient-rich Millets. In: Singh U, Praharaj C, Singh S, Singh N (eds) Biofortification of food crops. Springer, New Delhi, pp 409–420Google Scholar
  67. Skalicky A, Meyers A, Adams W, Yang Z, Cook JT, Frank DA (2006) Child food insecurity and iron deficiency anemia in low income infants and toddlers in the United States. Matern Child Health J 10(2):177–185Google Scholar
  68. Stangoulis J, Guild G (2014) Measuring trace micronutrient levels in crops. Biofortification progress briefs, August 2014, pp 27–28Google Scholar
  69. Stangoulis JCR, Huynh BL, Welch RM, Choi EY, Graham RD (2007) Quantitative trait loci for phytate in rice grain and their relationship with grain micronutrients content. Euphytica 154(3):289–294CrossRefGoogle Scholar
  70. Stein AJ, Meenakshi JV, Qaim M (2005) Analyzing the health benefits of biofortified staple crops by means of the disability-adjusted life years approach: a handbook focusing on iron, zinc and vitamin A. HarvestPlus technical monograph 4. IFPRI/CIAT, Washington, DCGoogle Scholar
  71. Tako E, Beebe SE, Reed S, Hart JJ, Glahn RP (2014) Polyphenolic compounds appear to limit the nutritional benefit of biofortified higher iron black bean (Phaseolus vulgaris L.). Nutr J 13:28Google Scholar
  72. Tako E, Reed SM, Budiman J, Hart JJ, Glahn RP (2015) Higher iron pearl millet (Pennisetum glaucum L.) provides more absorbable iron that is limited by increased polyphenolic content. Nutr J 14:11Google Scholar
  73. UNICEF (2004) Vitamin and mineral deficiency, a global progress report. UNICEF and micronutrient initiative.
  74. Velu G, Rai KN, Muralidharan V, Kulkarni VN, Longvah T, Raveendran TS (2007) Prospects of breeding biofortified pearl millet with high grain iron and zinc contents. Plant Breed 126(2):182–185CrossRefGoogle Scholar
  75. Velu G, Rai KN, Sahrawat KL et al (2008) Variability for grain iron and zinc contents in pearl millet hybrids. J SAT Agric Res 6:1–4Google Scholar
  76. Velu G, Rai KN, Muralidharan V (2011) Gene effects and heterosis for grain iron and zinc density in pearl millet (Pennisetum glaucum (L.) R. Br). Euphytica 180(2):251–259Google Scholar
  77. Vucenik I, Shamsuddin AM (2003) Cancer inhibition by inositol hexaphosphate (IP6) and inositol: from laboratory to clinic. J Nutr 133(11):3778S–3784SCrossRefGoogle Scholar
  78. Vucenik I, Shamsuddin AM (2006) Protection against cancer by dietary IP6 and inositol. Nutr Cancer 55(2):109–125CrossRefGoogle Scholar
  79. Waters BM, Grusak MA (2008) Quantitative trait locus mapping for seed mineral concentrations in two Arabidopsis thaliana recombinant inbred populations. New Phytol 179(4):1033–1047CrossRefGoogle Scholar
  80. Welch RM, Graham RD (1999) A new paradigm for world agriculture: meeting human needs: productive, sustainable, nutritious. Field Crop Res 60(1–2):1–10CrossRefGoogle Scholar
  81. White PJ, Broadley MR (2005) Biofortifying crops with essential mineral elements. Trends Plant Sci 10(12):586–593CrossRefGoogle Scholar
  82. White PJ, Broadley MR (2009) Biofortification of crops with seven mineral elements often lacking in human diets—iron, zinc, copper, calcium, magnesium, selenium and iodine. New Phytol 182:49–84CrossRefGoogle Scholar
  83. Winkler JT (2011) Biofortification: improving the nutritional quality of staple crops. In: Pasternak C (ed) Access not excess. Smith-Gordon Publishing, UK, pp 100–112Google Scholar
  84. Zern TL, Fernandez ML (2005) Cardioprotective effects of dietary polyphenols. J Nutr 135(10):2291–2294CrossRefGoogle Scholar

Copyright information

© Springer Nature Switzerland AG 2019

Authors and Affiliations

  1. 1.Department of BotanySt. Xavier’s College (Autonomous)PalayamkottaiIndia
  2. 2.T.A.L. Samy Unit for Plant Tissue Culture and Molecular Biology, Department of Plant Biology and BiotechnologyLoyola College (Autonomous)ChennaiIndia

Personalised recommendations