, Volume 240, Issue 6, pp 1167–1178 | Cite as

Wheat grain preharvest sprouting and late maturity alpha-amylase

  • Daryl J. Mares
  • Kolumbina Mrva


Preharvest sprouting (PHS) and late maturity α-amylase (LMA) are the two major causes of unacceptably high levels of α-amylase in ripe wheat grain. High α-amylase activity in harvested grain results in substantially lower prices for wheat growers and at least in the case of PHS, is associated with adverse effects on the quality of a range of end-products and loss of viability during storage. The high levels of α-amylase are reflected in low falling number, the internationally accepted measure for grain receival and trade. Given the significant losses that can occur, elimination of these defects remains a major focus for wheat breeding programs in many parts of the world. In addition, the genetic, biochemical and molecular mechanisms involved in the control of PHS and LMA as well as the interactions with environmental factors have attracted a sustained research interest. PHS and LMA are independent, genetically controlled traits that are strongly influenced by the environment, where the effects of particular environmental factors vary substantially depending on the stage of grain development and ripening. This review is a summary and an assessment of results of recent research on these important grain quality defects.


Dormancy Release from dormancy Abscisic acid Gibberellic acid Grain development QTL 



Preharvest sprouting


Late maturity α-amylase


Quantitative trait locus


Abscisic acid


Gibberellic acid


Polymerase chain reaction


Synthetic hexaploid wheat



The authors gratefully acknowledge funding support from the Australian Grains Research and Development Corporation and thank Dr Peter Chandler, CSIRO, Australia for helpful discussion of the possible role of gibberellins in LMA.

Conflict of interest

We declare that we have no conflict of interest.

Supplementary material

425_2014_2172_MOESM1_ESM.docx (17 kb)
Supplementary material 1 (DOCX 17 kb)
425_2014_2172_MOESM2_ESM.docx (18 kb)
Supplementary material 2 (DOCX 17 kb)


  1. Anderson JA, Sorrells ME, Tanksley SD (1993) RFLP analysis of genomic regions associated with resistance to preharvest sprouting in wheat. Crop Sci 33:453–459CrossRefGoogle Scholar
  2. Ashikawa I, Abe F, Nakamura S (2010) Ectopic expression of wheat and barley DOG1-like genes promotes seed dormancy in Arabidopsis. Plant Sci 179:536–542PubMedCrossRefGoogle Scholar
  3. Barrero JM, Talbot MJ, White RG, Jacobsen JV, Gubler F (2009) Anatomical and transcriptomic studies of the coleorhiza reveal the importance of this tissue in regulating dormancy in barley. Plant Physiol 150:1006–1021PubMedCentralPubMedCrossRefGoogle Scholar
  4. Barrero JM, Mrva K, Talbot MJ, White JT, Gubler F, Mares DJ (2013) Genetic, hormonal and physiological analysis of late maturity α-amylase (LMA) in wheat. Plant Physiol 161:1265–1277PubMedCentralPubMedCrossRefGoogle Scholar
  5. Benech-Arnold RL, Gualano N, Leymarie J, Come D, Corbineau F (2006) Hypoxia interferes with ABA metabolism and increases ABA sensitivity of dormant barley grains. J Exp Bot 57:1423–1430PubMedCrossRefGoogle Scholar
  6. Bewley D (1997) Seed germination and dormancy. Plant Cell 9:1055–1066PubMedCentralPubMedCrossRefGoogle Scholar
  7. Biddulph TB, Plummer JA, Setter TL, Mares DJ (2007) Influence of high temperature and terminal moisture stress on dormancy in wheat (Triticum aestivum L.). Field Crops Res 103:139–153CrossRefGoogle Scholar
  8. Canadian Grain Commission Fact Sheet “Sprouted and severely sprouted kernels”
  9. Chang C, Zhang H-P, Zhao Q-X, Feng J-M, Si H-Q, Lu J, Ma C-X (2011) Rich allelic variations of Viviparous-1A and their associations with seed dormancy/pre-harvest sprouting of common wheat. Euphytica 179:343–353CrossRefGoogle Scholar
  10. Chao S, Xu SS, Elias EM, Faris JD, Sorrells ME (2010) Identification of chromosome locations of genes affecting preharvest sprouting and seed dormancy using chromosome substitution lines in tetraploid wheat (Triticum turgidum L.). Crop Sci 50:1180–1187CrossRefGoogle Scholar
  11. Cheng C-R, Oldack K, Mrva K, Mares D (2014) Analysis of high pI α-Amy-1 gene family members expressed in late maturity a-amylase in wheat (Triticum aestivum L.). Mol Breed 33:519–529PubMedCentralPubMedCrossRefGoogle Scholar
  12. DePauw RM, McCaig TN (1991) Components of variation, heritabilities and correlations for indices of sprouting tolerance and seed dormancy in Triticum spp. Euphytica 52:221–229CrossRefGoogle Scholar
  13. Derera NF (1989a) Preharvest Sprouting in Cereals. CRC Press Inc., Boca RatonGoogle Scholar
  14. Derera NF (1989b) Breeding for preharvest sprouting tolerance. In: Derera NF (ed) Preharvest sprouting in cereals. CRC Press Inc., Boca Raton, pp 111–128Google Scholar
  15. Edwards MM (1973) Seed dormancy and environment: internal oxygen relationships. In: Heydecker W (ed) Seed ecology. Butterworths, London, UK, pp 169–188Google Scholar
  16. Edwards RA, Ross AS, Mares DJ, Ellison FW, Tomlinson JD (1989) Enzymes from rain-damaged and laboratory germinated wheat I. Effects on product quality. J Cereal Sci 10:157–167CrossRefGoogle Scholar
  17. Emebri LC, Oliver JR, Mrva K, Mares DJ (2010) Association mapping of late maturity a-amylase (LMA) activity in a collection of synthetic hexaploid wheat. Mol Breed 26:39–49CrossRefGoogle Scholar
  18. Fernandez MR, Sissons M, Conner RL, Wang H, Clarke JM (2011) Influence of biotic and abiotic factors on dark discoloration of durum wheat kernels. Crop Sci 51:1205–1214CrossRefGoogle Scholar
  19. Flintham JE (2000) Different genetic components control coat-imposed and embryo-imposed dormancy in wheat. Seed Sci Res 10:43–50CrossRefGoogle Scholar
  20. Flintham J, Adlam R, Gale M (1999) Eighth International Symposium on Pre-Harvest Sprouting in Cereals 1998. Weipert D (ed). Association of Cereal Research. Federal Centre for Cereal, Potato and Lipid Research, Detmold, Germany, pp 67–76Google Scholar
  21. Flintham JE, Adlam R, Bassoi M, Holdsworth M, Gale M (2002) Mapping genes for resistance to sprouting damage in wheat. Euphytica 126:39–45CrossRefGoogle Scholar
  22. Fofana B, Humphreys DG, Cloutier S, Brule-Babel A, Woods S, Luckow OM, Somers DJ (2009) Mapping quantitative trait loci controlling pre-harvest sprouting resistance in a red 3 white seeded spring wheat cross. Euphytica 165:509–521CrossRefGoogle Scholar
  23. Gaskin P, Kirkwood PS, Lenton JR, MacMIllan J, Radley ME (1980) Identification of gibberellins in developing wheat grain. Agric Biol Chem 44:1589–1593CrossRefGoogle Scholar
  24. Gerjets T, Scholefield D, Foulkes MJ, Lenton JR, Holdsworth MJ (2009) An analysis of dormancy, ABA responsiveness, after-ripening and pre-harvest sprouting in hexaploid wheat (Triticum aestivum L.) caryopses. J Exp Bot 61:597–607PubMedCentralPubMedCrossRefGoogle Scholar
  25. Himi E, Maekawa M, Miura H, Noda K (2011) Development of PCR markers for Tamyb10 related to R-1, red grain color gene in wheat. Theor Appl Genet 122:1561–1576PubMedCrossRefGoogle Scholar
  26. Huang T, Qu B, Li H-P, Zuo Z-X, Liao Y-C (2012) A maize viviparous 1 gene increases seed dormancy and preharvest sprouting tolerance in transgenic wheat. J Cereal Sci 55:166–173CrossRefGoogle Scholar
  27. Jacobsen JV, Barrero JM, Hughes T, Julkowska M, Taylor JM, Xu Q, Gubler F (2013) Roles for blue light, jasmonate and nitric oxide in the regulation of dormancy and germination in wheat grain (Triticum aestivum L.). Planta 238:121–138PubMedCrossRefGoogle Scholar
  28. Kato K, Nakamura W, Tabiki T, Miura H, Sawada S (2001) Detection of loci controlling seed dormancy on group 4 chromosomes of wheat and comparative mapping with rice and barley genomes. Theor Appl Genet 102:980–985CrossRefGoogle Scholar
  29. King RW (1989) Physiology of sprouting resistance. In: Derera NF (ed) Preharvest field sprouting in cereals. CRC Press Inc, Boca Ratan, pp 27–60Google Scholar
  30. Kondhare KR, Kettlewell PS, Farrell AD, Hedden P, Monaghan JM (2012) Effects of exogenous abscisic acid and gibberellic acid on pre-maturity α-amylase formation in wheat grains. Euphytica 181:51–60Google Scholar
  31. Kondhare KR, Kettlewell PS, Farrell AD, Hedden P, Monaghan JM (2013) The role of sensitivity to abscisic acid and gibberellin in pre-maturity α-amylase formation in wheat grains. J Cereal Sci 58:472–478CrossRefGoogle Scholar
  32. Kondhare KR, Hedden P, Kettlewell PS, Farrell AD, Monaghan JM (2014) Use of the hormone-biosynthesis inhibitors fluridone and paclobutrazol to determine the effects of altered abscisic acid and gibberellin levels on pre-maturity a-amylase formation in wheat grains. J Cereal Sci 60:210–216CrossRefGoogle Scholar
  33. Lenoir C, Corbineau F, Come D (1986) Barley (Hordeum vulgare) seed dormancy as related to glumella characteristics. Physiol Plant 68:301–307CrossRefGoogle Scholar
  34. Lenton JR, Gale MD (1987) Hormonal changes during cereal grain development. In: Mares DJ (ed) 4th international symposium on pre-harvest sprouting in cereals. Westview Press, Boulder, pp 253–264Google Scholar
  35. Liu S, Bai G (2010) Dissection and fine mapping of a major QTL for preharvest sprouting in white wheat Rio Blanco. Theor Appl Genet 121:1395–1404Google Scholar
  36. Liu S, Sehgal SK, Li J, Lin M, Trick HN, Yu J, Gill BS, Bai G (2013a) Cloning and characterization of a critical regulator for preharvest sprouting in wheat. Genetics 195:263–273PubMedCentralPubMedCrossRefGoogle Scholar
  37. Liu A, Gao F, Kanno Y, Jordan MC, Kamiya Y, Seo M, Ayele B (2013b) regulation of wheat seed dormancy by after-ripening is mediated by specific transcriptional switches that induce changes in seed hormone metabolism and signalling. PLoS One 8:e56570PubMedCentralPubMedCrossRefGoogle Scholar
  38. Mares DJ (1983) Investigation of the preharvest sprouting resistance mechanisms in some Australian wheats. In: Kruger JE, LaBerge DE (eds) Third international symposium on preharvest sprouting in cereals. Westview Press Inc., Boulder Co, pp 59–65Google Scholar
  39. Mares DJ (1984) Temperature dependence of germinability of wheat (Triticum aestivum L.) grain in relation to pre-harvest sprouting. Aust J Agric Res 35:115–128CrossRefGoogle Scholar
  40. Mares DJ (1987) Preharvest sprouting in white grained wheats. In: Mares DJ (ed) Fourth international symposium on preharvest sprouting in cereals. Westview Press Inc., Boulder Co, pp 64–74Google Scholar
  41. Mares DJ (1989) Preharvest sprouting damage and sprouting tolerance: Assay methods and instrumentation. In: Derera NF (ed) Preharvest sprouting in cereals. CRC Press Inc., Boca Raton, pp 129–170Google Scholar
  42. Mares DJ (1993) Genetic studies of sprouting tolerance in red and white wheats. In: Walker-Simmons MK, Ried JL (eds) Pre-Harvest Sprouting in Cereals 1992. American association of cereal chemists, St Paul, pp 21–29Google Scholar
  43. Mares DJ (1999) The seed coat and dormancy in wheat grains. In: Weipert D (ed) Eighth international symposium on pre-harvest sprouting in cereals 1998. Association of Cereal Research, Federal Centre for Cereal, Potato and Lipid Research, Detmold, pp 77–81Google Scholar
  44. Mares DJ, Mrva K (2001) Mapping quantitative trait loci associated with variation in grain dormancy in Australian wheat. Aust J Agric Res 52:1257–1265CrossRefGoogle Scholar
  45. Mares DJ, Mrva K (2008a) Genetic variation for quality traits in synthetic wheat germplasm. Aust J Agric Res 59:406–412CrossRefGoogle Scholar
  46. Mares D, Mrva K (2008b) Late-maturity a-amylase: low falling number in wheat in the absence of preharvest sprouting. J Cereal Sci 47:6–17CrossRefGoogle Scholar
  47. Mares DJ, Mrva K, Cheong J, Williams K, Watson B, Storlie E, Sutherland M, Zou Y (2005) A QTL located on chromosome 4A associated with dormancy in white- and red-grained wheats of diverse origin. Theor Appl Genet 111:1357–1364PubMedCrossRefGoogle Scholar
  48. Mares DJ, Rathjen J, Mrva K, and Cheong J (2009) Genetic and environmental control of dormancy in white-grained wheat (Triticum aestivum L.) Euphytica 168:311-318.Google Scholar
  49. Mares DJ, Rathjen J, Mrva K, Cheong J, Oldach K (2012). Germinability during grain development and after-ripening in wheat lines carrying different alleles at dormancy QTL. J Nyachiro (ed). In: 12th International Symposium on Preharvest Sprouting in Cereals, Canada 2012 AbstractGoogle Scholar
  50. McCarty DR, Carson CB, Stinard PS, Robertson DS (1989) Molecular analysis of vívíparous-I: an abscisic acid insensitive mutant of Maize. Plant Cell 1:523–532PubMedCentralPubMedCrossRefGoogle Scholar
  51. McKibbin RS, Wilkinson MD, Bailey PC, Flintham JE, Andrew LM, Lazzeri PA, Gale MD, Lenton JR, Holdsworth MJ (2002) Transcripts of Vp-1 homeologues are misspliced in modern wheat and ancestral species. PNAS 99:10203–10208PubMedCentralPubMedCrossRefGoogle Scholar
  52. Mohler V, Albrecht T, Mrva K, Schweizer G, Hartl L (2014) Genetic analysis of falling number in three bi-parental common winter wheat populations. Plant Breed 133:448–453CrossRefGoogle Scholar
  53. Mori M, Uchino N, Chono M, Kato K, Miura H (2005) Mapping QTLs for grain dormancy on wheat chromosome 3A and the group 4 chromosomes, and their combined effect. Theor Appl Genet 110:1315–1323PubMedCrossRefGoogle Scholar
  54. Mrva K, Mares DJ (1996a) Inheritance of late maturity α-amylase in wheat. Euphytica 88:61–67CrossRefGoogle Scholar
  55. Mrva K, Mares DJ (1996b) Expression of late maturity α-amylase in wheat containing gibberellic acid insensitivity genes. Euphytica 88:68–76Google Scholar
  56. Mrva K, Mares DJ (1999) Regulation of high pI α-amylase synthesis in wheat aleurone by a gene(s) located on chromosome 6B. Euphytica 109:17–23CrossRefGoogle Scholar
  57. Mrva K, Mares DJ (2001) Quantitative trait locus analysis of late maturity α-amylase in wheat using the doubled haploid population Cranbrook x Halberd. Aust J. Agric Res 52:1267–1273CrossRefGoogle Scholar
  58. Mrva K, Wallwork M, Mares DJ (2006) a-Amylase and programmed cell death in aleurone of ripening wheat grains. J Expt Botany 57:877–885CrossRefGoogle Scholar
  59. Mrva K, Mares DJ, Cheong J (2008) Genetic mechanisms involved in late maturity α-amylase in wheat. In: Appels R, Eastwood R, Lagudah E, Langridge P, MacKay M, McIntyre L, Sharp P (eds). Proceedings of the 11th International Wheat Genetics Symposium, Brisbane 2008. Sydney University Press pp 940–942Google Scholar
  60. Mrva K, Cheong J, Yu B, Law HY, Mares DJ (2009) Late maturity α-amylase in synthetic hexaploid wheat. Euphytica 168:403–411CrossRefGoogle Scholar
  61. MrvaK, Mares DJ(1996) Control of late maturity α-amylase synthesis compared to enzyme synthesis during germination. In: Noda K, Mares DJ (eds) Preharvest Sprouting in Cereals 1995. Centre for Academic Societies Osaka, pp 419–426Google Scholar
  62. Muncvold JD, Tanaka J, Benscher D, Sorrells ME (2009) Mapping quantitative trait loci for preharvest sprouting resistance in white wheat. Theor Appl Genet 119:1223–1235CrossRefGoogle Scholar
  63. Nakamura S, Abe F, Kawahigashi H, Nakazono K, Tagiri A, Matsumoto T, Utsugi S, Ogawa T, Handa H, Ishida H, Mori M, Kawaura K, Ogihara Y, Miura H (2011) A Wheat homolog of MOTHER OF FT AND TFL1 acts in the regulation of germination. Plant Cell 23:3215–3229PubMedCentralPubMedCrossRefGoogle Scholar
  64. Noda K, Kawabata C, Kawakami N (1993) Wheat grain imbibition at low temperatures and embryo responsiveness to ABA. In: Walker-Simmons MK, Ried JL (eds) Preharvest Sprouting in Cereals 1992. American Association of Cereal Chemists, St Paul, pp 367–372Google Scholar
  65. Noda K, Matsuura T, Maekawa M, Taketa S (2002) Chromosomes responsible for sensitivity of embryo to abscisic acid and dormancy in wheat. Euphytica 123:203–209CrossRefGoogle Scholar
  66. Noll JS, Dyck DL, Czarnecki E (1982) Expression of RL4137 type of dormancy in F1 seeds of reciprocal crosses in common wheat. Can J Plant Sci 62:345–349CrossRefGoogle Scholar
  67. Osa M, Kato K, Mori M, Shindo C, Torada A, Miura H (2003) Mapping QTLs for seed dormancy and the Vp1 homologue on chromosome 3A in wheat. Theor Appl Genet 106:1491–1496PubMedGoogle Scholar
  68. Rathjen JR, Strounina EV, Mares DJ (2009) Pathway for Water Movement into Dormant and Non-Dormant Wheat (Triticum aestivum L.) Grains. J Exp Bot 60:1619–1631PubMedCentralPubMedCrossRefGoogle Scholar
  69. Schramm EC, Nelson SK, Kidwell KK, Steber CM (2013) Increased ABA sensitivity results in higher seed dormancy in soft white spring wheat cultivar ‘Zak’. Theor Appl Genet 126:791–803PubMedCrossRefGoogle Scholar
  70. Somyong S, Munckvold JD, Tanaja J, Benscher D, Sorrells ME (2011) Comparative genetic analysis of a wheat seed dormancy QTL with rice and Brachypodium identifies candidate genes for ABA perception and calcium signalling. Funct Integr Genomics 11:479–490PubMedCrossRefGoogle Scholar
  71. Tomkinson CL, Lyndon RF, Arnold GM, Lenton JR (1997) The effects of temperature and the Rht3 dwarfing gene on growth, cell extension, and gibberellin content and responsiveness in the wheat leaf. J Exp Bot 48:963–970CrossRefGoogle Scholar
  72. Torada A, Koike M (2012) Marker assisted development and evaluation of near isogenic lines for seed dormancy genes on chromosome 3A and 4A in bread wheat. J Nyachiro (ed). In: 12th International Symposium on Preharvest Sprouting in Cereals, Canada 2012. AbstractGoogle Scholar
  73. Trethowan RM (1995) Evaluation and selection of bread wheat (Triticum aestivum L.) for preharvest sprouting tolerance. Aust J Agric Res 46:463–474CrossRefGoogle Scholar
  74. Verrity JCK, Hac L, Skerritt JH (1999) Development of a field enzyme-linked immunosorbent assay (ELISA) for detection of a-amylase in preharvest-sprouted wheat. Cereal Chem 76:673–681CrossRefGoogle Scholar
  75. Walker-Simmons MK (1987) ABA levels and sensitivity in developing wheat embryos of sprouting resistant and susceptible cultivars. Plant Physiol 84:61–66PubMedCentralPubMedCrossRefGoogle Scholar
  76. Warner RL, Kudrna DA, Spaeth SC, Jones SS (2000) Dormancy in white-grain mutants of Chinese Spring wheat (Triticum aestivum L.). Seed Sci Res 10:51–60Google Scholar
  77. Williamson PM (1997) Black point of wheat: in vitro production of symptoms, enzymes involved, and association with Alternaria alternate. Aust J Agric Res 48:13–19CrossRefGoogle Scholar
  78. Wilson ID, Barker GLA, Lu C, Coghill JA, Beswick RW, Lenton JR, Edwards KJ (2005) Alteration of the embryo transcriptome of hexaploid winter wheat (Triticum aestivum cv. Mercia) during maturation and germination. Funct Integr Genomics 5:144–154Google Scholar
  79. Yang Y, Zhao XL, Xia LQ, Chen XM, Xia XC, Yu Z, He H, Roder M (2007) Development and validation of a Viviparous-1 STS marker for pre-harvest sprouting tolerance in Chinese wheats. Theor Appl Genet 115:971–980PubMedCrossRefGoogle Scholar
  80. Yang Y, Zhang CL, Liu SX, Sun YQ, Meng JY, Xia LQ (2014) Characterization of the rich haplotypes of Viviparous-1A in Chinese wheats and development of a novel sequence tagged site marker for pre-harvest sprouting resistance. Mol Breed 33:75–88CrossRefGoogle Scholar
  81. Zhang Y, Miao X, Xia X, He Z (2014) Cloning of seed dormancy genes (TaSdr) associated with toleranceto pre-harvest sprouting in common wheat and development of a functional marker. Theor Appl Genet 127:855–866PubMedCrossRefGoogle Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2014

Authors and Affiliations

  1. 1.School of Agriculture Food and WineUniversity of Adelaide, Waite CampusGlen OsmondAustralia

Personalised recommendations