Plant and Soil

, Volume 411, Issue 1–2, pp 69–79 | Cite as

Differences in grain zinc are not correlated with root uptake and grain translocation of zinc in wild emmer and durum wheat genotypes

  • Ozlem Yilmaz
  • Gamze Altintas Kazar
  • Ismail Cakmak
  • Levent Ozturk
Regular Article


Background and aims

Cereal-based foods fall short of providing adequate dietary zinc (Zn) to human beings. Developing new genotypes with high genetic capacity for root uptake and grain deposition of Zn is an important challenge. There is a large genetic variation for grain Zn concentration among and between wheat species, especially within wild emmer wheat (Triticum turgidum ssp. dicoccoides) that can be exploited in order to understand the physiological mechanisms contributing to grain Zn accumulation.


Eight different wild emmer genotypes and two durum wheat (Triticum durum) cultivars were used to investigate root uptake, root-to-shoot translocation and remobilization (i.e., retranslocation) from flag leaves into grains of 65ZnSO4-treated plants. The initial seed Zn concentrations of wild emmer wheat and durum genotypes used in the experiments were different, ranging from 45 to 73 mg kg−1 and from 35 to 40 mg kg−1, respectively. Plants were grown in nutrient solution for the experiments investigating root uptake and shoot transport of Zn by using 65Zn labeled ZnSO4 and in soil medium for the experiments studying shoot and grain Zn concentrations and 65Zn translocation from flag leaves into grains. The treatment of flag leaves with 65Zn was realized by immersion of flag leaves into 65ZnSO4 solution for 15 seconds and for 5 times during the anthesis and early milk stages.


Wild emmer and durum wheat genotypes expressed highly significant differences in root uptake and root-to-shoot translocation of 65Zn and translocation of 65Zn from flag leaves into grains. However, none of these parameters showed a significant correlation either with the initial seed Zn concentrations at sowing or the grain Zn concentrations at harvest. The durum wheat cultivars with higher grain yield had lower concentration of Zn both in seeds at sowing or in grains at harvest, while wild emmer genotypes with lower grain yield capacity had higher concentration of Zn both in seeds at sowing or in grains at harvest. The concentration or content (total amount) of Zn in shoot during the early growth stage also did not correlate with the initial seed Zn concentrations.


Differences in grain Zn concentration of wild emmer and cultivated wheats could not be explained by root Zn uptake and Zn translocation from flag leaf into grains during seedling and reproductive growth stages, respectively. It seems that there are additional key factors affecting the expression of genetic variation for grain Zn accumulation.


Zinc uptake Zinc translocation Grain zinc Wild emmer wheat Durum wheat 



The authors acknowledge TUBITAK (The Scientific and Technological Research Council of Turkey, project no. 108 T436) and the HarvestPlus Challenge Program for their financial support, Prof. Dr. Hakan Ozkan of Cukurova University for providing the seed material and Dr. Stuart James Lucas of Sabanci University for proof-reading of the manuscript.

Supplementary material

11104_2016_2969_MOESM1_ESM.docx (12 kb)
ESM 1 Table S1 (DOCX 11 kb)


  1. Bashir K, Takahashi R, Nakanishi H, Nishizawa NK (2013) The road to micronutrient biofortification of rice: progress and prospects. Frontiers Plant Sci 4:1–7CrossRefGoogle Scholar
  2. Bouis HE, Welch RM (2009) Biofortification—a sustainable agricultural strategy for reducing micronutrient malnutrition in the global south. Crop Sci 50:20–32CrossRefGoogle Scholar
  3. Cakmak I (2008) Enrichment of cereal grains with zinc: agronomic or genetic biofortification? Plant Soil 302:1–17CrossRefGoogle Scholar
  4. Cakmak I, Ekiz H, Yılmaz A, Torun B, Köleli N, Gültekin I, Alkan A, Eker S (1997) Differential response of rye, triticale, bread and durum wheats to zinc deficiency in calcareous soils. Plant Soil 188:1–10CrossRefGoogle Scholar
  5. Cakmak I, Torun B, Erenoglu B, Oztürk L, Marschner H, Kalaycı M, Ekiz H, Yılmaz A (1998) Morphological and physiological differences in cereals in response to zinc deficiency. Euphytica 100:349–357CrossRefGoogle Scholar
  6. Cakmak I, Tolay I, Ozdemir A, Ozkan H, Kling CI (1999) Differences in zinc efficiency among and within diploid, tetraploid and hexaploid wheats. Ann Bot 84:163–171CrossRefGoogle Scholar
  7. Cakmak I, Torun A, Millet E, Feldman M, Fahima T, Korol A, Nevo E, Braun HJ, Ozkan H (2004) Triticum dicoccoides: an important genetic resource for increasing zinc and iron concentration in modern cultivated wheat. Soil Sci Plant Nutr 50:1047–1054CrossRefGoogle Scholar
  8. Cakmak I, Pfeiffer WH, McClafferty B (2010a) Biofortification of durum wheat with zinc and iron. Cereal Chem 87:10–20CrossRefGoogle Scholar
  9. Cakmak I, Kalayci M, Kaya Y, Torun AA, Aydın N, Wang Y, Arısoy Z, Erdem H, Yazici A, Gokmen O, Ozturk L, Horst WJ (2010b) Biofortification and localization of zinc in wheat grain. J Agric Food Chem 58:9092–9102CrossRefPubMedGoogle Scholar
  10. Distelfeld A, Cakmak I, Peleg Z, Ozturk L, Yazici MA, Budak H, Saranga Y, Fahima T (2007) Multiple QTL-effects of wheat GpcB1 locus on grain protein and micronutrient concentrations. Physiol Plant 129:635–643CrossRefGoogle Scholar
  11. Edgerton MD (2009) Increasing crop productivity to meet global needs for feed, food, and fuel. Plant Physiol 149:7–13CrossRefPubMedPubMedCentralGoogle Scholar
  12. Fan MS, Zhao FJ, Fairweather-Tait SJ, Poulton PR, Dunham SJ, McGrath SP (2008) Evidence of decreasing mineral density in wheat grain over the last 160 years. J Trace Elem Med Biol 22:315–324CrossRefPubMedGoogle Scholar
  13. FAOSTAT (2011) Food and Agriculture Organization of the United Nations, FAOSTAT database, available at
  14. Gomez-Becerra HF, Erdem H, Yazici A, Tutus Y, Torun B, Ozturk L, Cakmak I (2010) Grain concentrations of protein and mineral nutrients in a large collection of spelt wheat grown under different environments. J Cereal Sci 52:342–349CrossRefGoogle Scholar
  15. Grassini P, Eskridge KM, Cassman KG (2013) Distinguishing between yield advances and yield plateaus in historical crop production trends. Nat Commun 4:2918. doi: 10.1038/ncomms3918 CrossRefPubMedPubMedCentralGoogle Scholar
  16. Harris D, Rashid A, Miraj G, Arif M, Shah H (2007) ‘on-farm’ seed priming with zinc sulphate solution—a cost-effective way to increase the maize yields of resource-poor farmers. Field Crops Res 102:119–127CrossRefGoogle Scholar
  17. Hussain S, Rengel Z, Mohammadi SA, Ebadi-Segherloo A, Maqsood MA (2016) Mapping QTL associated with remobilization of zinc from vegetative tissues into grains of barley (Hordeum vulgare). Plant Soil 399:193–208CrossRefGoogle Scholar
  18. Jiang W, Struik PC, van KH, Zhao M, Jin LN, Stomph TJ (2008) Does increased Zn uptake enhance grain Zn mass concentration in rice? Ann Appl Biol 153:135–147CrossRefGoogle Scholar
  19. Kutman UB, Yildiz BK, Ceylan Y, Ova EA, Cakmak I (2012) Contributions of root uptake and remobilization to grain zinc accumulation in wheat depending on post-anthesis zinc availability and nitrogen nutrition. Plant Soil 361:177–187CrossRefGoogle Scholar
  20. Lindsay WL, Norvell WA (1978) Development of a DTPA soil test for zinc, iron, manganese, and copper. Soil Sci Soc Am J 42:421–428CrossRefGoogle Scholar
  21. Lönnerdal B (2000) Dietary factors influencing zinc absorption. J Nutr 130:1378–1383Google Scholar
  22. Mabesa RL, Impa SM, Grewal S, Johnson-Beebout SE (2013) Contrasting grain-Zn response of biofortification rice (Oryza sativa L.) breeding lines to foliar Zn application. Field Crops Res 149:223–233CrossRefGoogle Scholar
  23. McDonald GK, Genc Y, Graham RD (2008) A simple method to evaluate genetic variation in grain zinc concentration by correcting for differences in grain yield. Plant Soil 306:49–55CrossRefGoogle Scholar
  24. Morgounov A, Belan I, Zelenskiy Y, Roseeva L, Tömösközi S, Békés F, Abugalieva A, Cakmak I, Vargas M, Crossa J (2013) Historical changes in grain yield and quality of spring wheat varieties cultivated in Siberia from 1900 to 2010. Can J Plant Sci 93:425–433CrossRefGoogle Scholar
  25. Murphy KM, Reeves PG, Jones SS (2008) Relationship between yield and mineral nutrient concentrations in historical and modern spring wheat cultivars. Euphytica 163:381–390CrossRefGoogle Scholar
  26. Olsen LI, Palmgren MG (2014) Many rivers to cross: the journey of zinc from soil to seed. Front Plant Sci 5:1–6Google Scholar
  27. Ortiz-Monasterio I, Graham RD (2000) Breeding for trace mineral in wheat. Food Nutr Bull 21:392–396CrossRefGoogle Scholar
  28. Pearson JN, Rengel Z (1994) Distribution and remobilization of Zn and Mn during grain development in wheat. J Exp Bot 45:1829–1835CrossRefGoogle Scholar
  29. Peleg Z, Saranga Y, Yazici MA, Fahima T, Ozturk L, Cakmak I (2008) Grain zinc, iron and protein concentrations and zinc-efficiency in wild emmer wheat under contrasting irrigation regimes. Plant Soil 306:57–67CrossRefGoogle Scholar
  30. Peleg Z, Cakmak I, Ozturk L, Yazici A, JunY BH, Korol AB, Fahima T, Saranga Y (2009) Quantitative trait loci conferring grain mineral nutrient concentrations in durum wheat x wild emmer wheat RIL population. Theor App Gen 119:353–369CrossRefGoogle Scholar
  31. Pfeiffer WH, McClafferty B (2007) Harvest plus: breeding crops for better nutrition. Crop Sci 47:88–105CrossRefGoogle Scholar
  32. Phattarakul N, Rerkasem B, Li LJ, Wu LH, Zou CQ, Ram H, Sohu VS, Kang BS, Surek H, Kalayci M, Yazici A, Zhang FS, Cakmak I (2012) Biofortification of rice grain with zinc through zinc fertilization in different countries. Plant Soil 361:131–141CrossRefGoogle Scholar
  33. Pottier M, Masclaux_Daubresse C, Yoshimoto K, Thomine S (2014) Autophagy as a possible mechanism for micronutrient remobilization from leaves to seeds. Frontiers in Plant Science 5:11CrossRefPubMedPubMedCentralGoogle Scholar
  34. Rengel Z, Batten GD, Crowley DE (1999) Agronomic approaches for improving the micronutrient density in edible portions of field crops. Field Crops Res 60:27–40CrossRefGoogle Scholar
  35. Sperotto RA (2013) Zn/Fe remobilization from vegetative tissues to rice seeds: should I stay or should I go? Ask Zn/Fe supply! Frontiers in Plant Sci 4:1–4CrossRefGoogle Scholar
  36. Sperotto RA, Ricachenevsky FK, Waldow VA, Müller ALH, Dressler VL, Fett JP (2013) Rice grain Fe, Mn and Zn accumulation: how important are flag leaves and seed number? Plant Soil Environ 59:262–266CrossRefGoogle Scholar
  37. Srinivasa J, Arun B, Mishra VK, Singh GP, Velu G, Babu R, Vasistha NK, Joshi AK (2014) Zinc and iron concentration QTL mapped in a Triticum spelta × T. aestivum cross. Theor Appl Genet 127:1643–1651CrossRefPubMedGoogle Scholar
  38. Stomph TJ, Jiang W, Struik PC (2009) Zinc biofortification of cereals: rice differs from wheat and barley. Trends Plant Sci 14:123–124CrossRefGoogle Scholar
  39. Tiwari C, Wallwork H, Arun B, Mishra VK, Velu G, Stangoulis JCR, Kumar U, Joshi AK (2016) Molecular mapping of quantitative trait loci for zinc, iron and protein content in the grains of hexaploid wheat. Euphytica 207:563–570CrossRefGoogle Scholar
  40. Uauy C, Distelfeld A, Fahima T, Blechl A, Dubcovsky J (2006) A NAC Gene regulating senescence improves grain protein, zinc, and iron content in wheat. Science 314:1298–1301CrossRefPubMedPubMedCentralGoogle Scholar
  41. Velu G, Ortiz-Monasterio I, Cakmak I, Hao Y, Singh RP (2014) Biofortification strategies to increase grain zinc and iron concentrations in wheat. J Cereal Sci 59:365–372CrossRefGoogle Scholar
  42. Von Wiren N, Marschner H, Romheld V (1996) Roots of iron-efficient maize also absorb phytosiderophore-chelated zinc. Plant Physiol 111:1119–1125CrossRefGoogle Scholar
  43. Waters BM, Sankaran RP (2011) Moving micronutrients from the soil to the seeds: genes and physiological processes from a biofortification perspective. Plant Sci 180:562–574CrossRefPubMedGoogle Scholar
  44. Waters BM, Uauy C, Dubcovsky J, Grusak MA (2009) Wheat (Triticum aestivum) NAM proteins regulate the translocation of iron, zinc, and nitrogen compounds from vegetative tissues to grain. J Exp Bot 60:4263–4274CrossRefPubMedGoogle Scholar
  45. Welch RM, House WA (1982) Availability to rats of zinc from soybean seeds as affected by maturity of seed, source of dietary-protein, and soluble phytate. J Nutr 112:879–885PubMedGoogle Scholar
  46. White PJ, Broadley MR (2005) Biofortifying crops with essential mineral elements. Trends Plant Sci 10:586–583CrossRefPubMedGoogle Scholar
  47. Xu Y, Ana D, Liub D, Zhang A, Xua H, Li B (2012) Molecular mapping of QTLs for grain zinc, iron and protein concentration of wheat across two environments. Field Crop Res 138:57–62CrossRefGoogle Scholar
  48. Zadoks JC, Chang TT, Konzak CF (1974) A decimal code for the growth stages of cereals. Weed Res 14:415–421CrossRefGoogle Scholar
  49. Zhang Y-Q, Sun YX, Ye Y-L, Karim MR, Xue Y-F, Yan P, Meng Q-F, Cui Z-L, Cakmak I, Zhang F-S, Zou C-Q (2011) Zinc biofortification of wheat through fertilizer applications in different locations of China. Field Crops Res 125:1–7CrossRefGoogle Scholar
  50. Zhao FJ, Su YH, Dunham SJ, Rakszegi M, Bedo Z, McGrath SP, Shewry PR (2009) Variation in mineral micronutrient concentrations in grain of wheat lines of diverse origin. J Cereal Sci 49:290–295CrossRefGoogle Scholar

Copyright information

© Springer International Publishing Switzerland 2016

Authors and Affiliations

  1. 1.Faculty of Science, Department of BiologyEge UniversityIzmirTurkey
  2. 2.Faculty of Science, Department of BiologyTrakya UniversityEdirneTurkey
  3. 3.Faculty of Engineering and Natural SciencesSabanci UniversityIstanbulTurkey

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