Uptake, distribution, and remobilization of iron and zinc among various tissues of wheat–Aegilops substitution lines at different growth stages

  • Prachi Sharma
  • Imran Sheikh
  • Dharmendra Singh
  • Satish Kumar
  • Shailender Kumar Verma
  • Rahul Kumar
  • Pritesh Vyas
  • Harcharan Singh Dhaliwal
Original Article


Biofortification of wheat for higher grain iron and zinc is the most feasible and cost-effective approach for alleviating micronutrient deficiency. The non-progenitor donor Aegilops species had 2–3 times higher grain iron and zinc content than the wheat cultivars, whereas the wheat–Aegilops substitution lines mostly of group 2 and 7 chromosomes had intermediate levels of grain micronutrients. The non-progenitor Aegilops species also had the highest iron content and intermediate-to-highest zinc content in straw, lower leaves, and flag leaves at the pre-anthesis, grain-filling, and maturity growth stages. The micronutrients accumulation status is followed by wheat–Aegilops substitution lines and is the least in wheat cultivars indicating that the donor Aegilops species and their substituted chromosomes possess genes for higher iron and zinc uptake and mobilization. The grain iron content was highly positively correlated with iron content in the plant tissues. Most of the lines had much higher iron and zinc content in all tissues during grain-filling period indicating higher iron and zinc uptake from soil during this stage. Although iron and zinc contents are nearly similar in grains, there was much less zinc content in the plant tissues of all the lines suggesting that the Triticeae species take up less zinc which is mobilized to grains more effectively than iron.


Iron and zinc Wheat–Aegilops substitution lines Biofortification and metal homeostasis genes Grain filling Flag leaves 



The authors acknowledge the Department of Biotechnology, Government of India for Grant (BT/AGR/Wheat Bioforti/PH-II/2010) through a network project “Biofortification of wheat for enhanced iron and zinc content by conventional and molecular breeding-Phase II”. The authors also acknowledge the Akal College of Agriculture for providing infrastructural facilities to carry out this work.

Supplementary material

11738_2017_2456_MOESM1_ESM.docx (16 kb)
Supplementary material 1 (DOCX 17 kb)


  1. Borrill P, Connorton JM, Balk J, Miller AJ, Sanders D, Uauy C (2014) Biofortification of wheat grain with iron and zinc: integrating novel genomic resources and knowledge from model crops. Front Plant Sci 5:53CrossRefPubMedPubMedCentralGoogle Scholar
  2. Bouis HE, Welch RM (2010) Bio-fortification: a sustainable agricultural strategy for reducing micronutrient malnutrition in the global south. Crop Sci 50:S20–S32CrossRefGoogle Scholar
  3. Cakmak I, Pfeiffer WH, McClafferty B (2010) Biofortification of durum wheat with zinc and iron. Cereal Chem 87:10–20CrossRefGoogle Scholar
  4. Farkas A, Molnar I, Dulai S, Rapi S, Oldal V, Cseh A, Kruppa K, Lang MM (2014) Increased micronutrient content (Zn, Mn) in the 3 Mb (4B) wheat-Aegilops biuncialis substitution and 3 Mb.4BS translocation identified by GISH and FISH. Genome 57:61–67CrossRefPubMedGoogle Scholar
  5. Genc Y, Verbyla AP, Torun AA, Cakmak I, Willsmore K, Wallwork H, McDonald GK (2009) Quantitative trait loci analysis of zinc efficiency and grain zinc concentration in wheat using whole genome average interval mapping. Plant Soil 314:49–66CrossRefGoogle Scholar
  6. Grillet L, Mari S, Schmidt W (2014) Iron in seeds—loading pathways and subcellular localization. Front Plant Sci 4:535CrossRefPubMedPubMedCentralGoogle Scholar
  7. Hanbidge KM (1987) Zinc. In: Mertz W (ed) Trace elements in human and animal nutrition, vol 5. Academic Press, Orlando, p 1Google Scholar
  8. Iwai T, Takahashi M, Oda K, Terada Y, Yoshida KT (2012) Dynamic changes in the distribution of minerals in relation to phytic acid accumulation during rice seed development. Plant Physiol 160:2007–2014CrossRefPubMedPubMedCentralGoogle Scholar
  9. Jorhem L, Engman J (2000) Determination of lead, cadmium, zinc, copper, and iron in foods by atomic absorption spectrometry after microwave digestion: NMKL collaborative study. J AOAC Int 83:1189–1203PubMedGoogle Scholar
  10. Kabata-Pendias A (2001) Trace elements in soils and plants. CRC Press, Boca Raton, p 140Google Scholar
  11. Klatte M, Schuler M, Wirtz M, Fink-Straube C, Hell R, Bauer P (2009) The analysis of Arabidopsis nicotianamine synthase mutants reveals functions for nicotianamine in seed iron loading and iron deficiency responses. Plant Physiol 150:257–271CrossRefPubMedPubMedCentralGoogle Scholar
  12. Kobayashi T, Nishizawa NK (2012) Iron uptake, translocation, and regulation in higher plants. Annu Rev Plant Biol 63:131–152CrossRefPubMedGoogle Scholar
  13. Kumari N, Rawat N, Tiwari V, Kumar S, Chhuneja P, Singh K, Randhawa GS, Dhaliwal HS (2011) Introgression of group 4 and 7 chromosomes of Ae. peregrina in wheat enhances grain iron and zinc density. Mol Breed 28:623–634CrossRefGoogle Scholar
  14. Masuda H, Ishimaru Y, Aung MS, Kobayashi T, Kakei Y, Takahashi M, Higuchi K, Nakanishi H, Nishizawa NK (2012) Iron biofortification in rice by the introduction of multiple genes involved in iron nutrition. Sci Rep 2:543CrossRefPubMedPubMedCentralGoogle Scholar
  15. Mayer JE, Pfeiffer WH, Beyer P (2008) Biofortified crops to alleviate micronutrient malnutrition. Curr Opin Plant Biol 11(2):166–170CrossRefPubMedGoogle Scholar
  16. Murray MG, Thomson WF (1980) Rapid isolation of high molecular weight plant DNA. Nucleic Acids Res 8:4321–4325CrossRefPubMedPubMedCentralGoogle Scholar
  17. Norton GJ, Deacon CM, Xiong L, Huang S, Meharg AA, Price AH (2010) Genetic mapping of the rice ionome in leaves and grain: Identification of QTLs for 17 elements including arsenic, cadmium, iron and selenium. Plant Soil 329:139–153CrossRefGoogle Scholar
  18. O’Brien TP, Summut ME, Lee JW, Smart MG (1985) The vascular system of the wheat spikelet. Aust J Plant Physiol 12:487–511CrossRefGoogle Scholar
  19. Olsen LI, Palmgren MG (2014) Many rivers to cross: the journey of zinc from soil to seed. Front Plant Sci 5:30PubMedPubMedCentralGoogle Scholar
  20. Palmgren MG, Clemens S, Williams LE, Kraemer U, Borg S, Schjorring JK, Sanders D (2008) Zinc biofortification of cereals: problems and solutions. Trends Plant Sci 13:464–473CrossRefPubMedGoogle Scholar
  21. Patrick JW, Offler CE (2001) Compartmentation of transport and transfer events in developing seeds. J Exp Bot 52:551–564CrossRefPubMedGoogle Scholar
  22. Peleg Z, Saranga Y, Yazici A, Fahima ZPT (2008) Grain zinc, iron and protein concentrations and zinc-efficiency in wild emmer wheat under contrasting irrigation regimes. Plant Soil 306:57–67CrossRefGoogle Scholar
  23. Peleg Z, Cakmak I, Ozturk L, Yazici A, Jun Y, Budak H, Korol AB, Fahima T, Saranga Y (2009) Quantitative trait loci conferring grain mineral nutrient concentrations in durum wheat × wild emmer wheat RIL population. Theor Appl Genet 119:353–369CrossRefPubMedGoogle Scholar
  24. Pfeiffer WH, McClafferty B (2007) HarvestPlus: breeding crops for better nutrition. Crop Sci 6:S88–S105Google Scholar
  25. Rawat N, Tiwari VK, Singh N, Randhawa GS, Singh K, Chhuneja P, Dhaliwal HS (2009) Evaluation and utilization of Aegilops and wild Triticum species for enhancing iron and zinc content in wheat. Genet Resour Crop Evol 56(1):53–64CrossRefGoogle Scholar
  26. Rawat N, Tiwari VK, Neelam K, Randhawa GS, Friebe B, Gill BS, Dhaliwal HS (2011) Development and molecular characterization of wheat-Aegilops kotschyi addition and substitution lines with high grain protein, iron and zinc. Genome 54:943–953CrossRefPubMedGoogle Scholar
  27. Sharma P, Imran Sharma P, Chugh V, Dhaliwal HS, Singh D (2014) Morphological, cytological and biochemical characterization of wheat Aegilops longissima derivatives BC1F6 and BC2F4 with high grain micronutrient. Int J Agric Environ Biotech 7(2):191–204CrossRefGoogle Scholar
  28. Tiwari VK, Rawat N, Chhuneja P, Neelam K, Aggarwal R, Randhawa GS, Dhaliwal HS, Keller B, Singh K (2009) Mapping of quantitative trait loci for grain iron and zinc concentration in diploid A genome wheat. J Hered 100:771–776CrossRefPubMedGoogle Scholar
  29. Tiwari VK, Rawat N, Neelam K, Kumar S, Randhawa GS, Dhaliwal HS (2010) Substitutions of 2S and 7U chromosomes of Aegilops kotschyi in wheat enhance grain iron and zinc concentration. Theor Appl Genet 121:259–269CrossRefPubMedGoogle Scholar
  30. Vasconcelos M, Datta K, Oliva N, Khalekuzzaman M, Torrizo L, Krishnan S, Oliveira M, Goto F, Datta SK (2003) Enhanced iron and zinc accumulation in transgenic rice with the ferritin gene. Plant Sci 164:371–378CrossRefGoogle Scholar
  31. Verma SK, Kumar S, Sheikh I, Malik S, Mathpal P, Chugh V, Kumar S, Prasad R, Dhaliwal HS (2016a) Transfer of useful variability of high grain iron and zinc from Aegilops kotschyi into wheat through seed irradiation approach. Int J Radiat Biol 92:132–139CrossRefPubMedGoogle Scholar
  32. Verma SK, Kumar S, Sheikh I, Sharma P, Mathpal P, Malik S, Kundu P, Awasthi A, Kumar S, Prasad R, Dhaliwal HS (2016b) Induced homoeologous pairing for transfer of useful variability for high grain Fe and Zn from Aegilops kotschyi into wheat. Plant Mol Biol Rep. doi: 10.1007/s11105-016-0989-8 Google Scholar
  33. Wada T, Lott JNA (1997) Light and electron microscopic and energy dispersive X-ray microanalysis studies of globoids in protein bodies of embryo tissues and the aleurone layer of rice (Oryza sativa L.) grains. Can J Bot 75:1137–1147CrossRefGoogle Scholar
  34. Waters BN, Sankaran RP (2011) Moving micronutrients from the soil to the seeds: genes and physiological processes from the biofortification perspectives. Plant Sci 180:562–574CrossRefPubMedGoogle Scholar
  35. Wei H, Fu Y, Arora R (2005) Intron-flanking EST-PCR markers: from genetic marker development to gene structure analysis in Rhododendron. Theor Appl Genet 111(7):1347–1356CrossRefPubMedGoogle Scholar
  36. Wirth J, Poletti S, Aeschlimann B, Yakandawala N, Drosse B, Osorio S, Tohge T, Fernie AR, Günther D, Gruissem W, Sautter C (2009) Rice endosperm iron biofortification by targeted and synergistic action of nicotianamine synthase and ferritin. Plant Biotechnol J 7(7):631–644CrossRefPubMedGoogle Scholar
  37. Zeidan MS, Mohamed MF, Hamouda HA (2010) Effect of foliar fertilization of Fe, Mn and Zn on wheat yield and quality in low sandy soils fertility. World J Agric Sci 6(6):696–699Google Scholar

Copyright information

© Franciszek Górski Institute of Plant Physiology, Polish Academy of Sciences, Kraków 2017

Authors and Affiliations

  • Prachi Sharma
    • 1
  • Imran Sheikh
    • 1
  • Dharmendra Singh
    • 2
  • Satish Kumar
    • 3
  • Shailender Kumar Verma
    • 4
  • Rahul Kumar
    • 1
  • Pritesh Vyas
    • 1
  • Harcharan Singh Dhaliwal
    • 1
  1. 1.Department of Biotechnology, Akal College of AgricultureEternal UniversityBaru SahibIndia
  2. 2.QAAFI, Centre of Plant SciencesThe University of QueenslandBrisbaneAustralia
  3. 3.Department of BiotechnologyIndian Institute of TechnologyRoorkeeIndia
  4. 4.School of Life SciencesCentral University of Himachal PradeshKangraIndia

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