Theoretical and Applied Genetics

, Volume 126, Issue 11, pp 2753–2762 | Cite as

An alternative mechanism for cleistogamy in barley

  • Ning Wang
  • Shunzong Ning
  • Mohammad Pourkheirandish
  • Ichiro Honda
  • Takao KomatsudaEmail author
Original Paper


Cleistogamy in barley is genetically determined by the presence of the recessive allele cly1, but the dominant allele at the linked locus Cly2 is epistatic over cly1. Although the molecular basis for cly1 action is well understood, that of Cly2 is not. Here we show that anther non-extrusion can occur not just when the lodicules fail to expand adequately (a trait which is fully determined by the allelic state at the cly1 locus), but by the premature timing of anthesis before the spike has emerged from the boot. The transcription of HvAP2 at cly1 is unaffected by the timing of anthesis. Where this occurs prematurely, by the time that the spike has emerged from the boot, the lodicules have already become shrunken and have lost the capacity to push the lemma and palea apart. Premature anthesis appears to be governed by a dominant gene, probably Cly2. Of the three phases of development of a non-cleistogamous barley floret (spike emergence from the boot, floret gaping induced by lodicule expansion and anther extrusion), genetic variation is available regarding at least the former two.


Leaf Sheath Anther Extrusion Flag Leaf Sheath Spike Emergence Premature Timing 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.



We thank K. Kakeda for useful comments on the manuscript. This research was funded by the Japanese Ministry of Agriculture, Forestry and Fisheries (Genomics for Agricultural Innovation grants no. TRG1004 and Genomics-based Technology for Agricultural Improvement grants no. TRS1002) to T.K. and the Japanese Society for the Promotion of Science (Postdoctoral Fellowship for Foreign Researchers) to N.W.

Supplementary material

122_2013_2169_MOESM1_ESM.pptx (379 kb)
Supplementary File 1 Spike emergence distance at the anthesis. (A) Measurement of spike emergence distance at the anthesis in F1 plant of SN × MG cross (left) and F1 plant of SN × KNG cross (right). (B) Frequency distribution of spike emergence distance in F2 population of SN × KNG cross. Genotypes for HvAP2/NmuCI are shown in different colors: black bars SN genotype (n = 32), light gray bars F1 genotype (n = 43), and white bars KNG genotype (n = 19). (C) F2 population of SN × MG cross: black bars SN genotype (n = 26), light gray bars F1 genotype (n = 49), and white bars MG genotype (n = 17). (D) F2 population of RIL50 × GP cross: black bars RIL50 genotype (n = 13), light gray bars F1 genotype (n = 30), and white bars GP genotype (n = 19) (PPTX 378 kb)
122_2013_2169_MOESM2_ESM.xls (20 kb)
Supplementary File 2 Spearman correlations between pairs of traits related to cleistogamy measured in the F2 population bred from the cross RIL50 × GP (n = 62)(XLS 20 kb)
122_2013_2169_MOESM3_ESM.xlsx (11 kb)
Supplementary File 3 Hypothetical genotypes contribute to lodicule size and spike emergence, and their overlapping effect on anther extrusion(XLSX 11 kb)
122_2013_2169_MOESM4_ESM.xls (30 kb)
Supplementary File 4 The sequences of PCR primers targeting the cly1 region(XLS 29 kb)


  1. Abdel-Ghani AH, Parzies HK, Omary A, Geiger HH (2004) Estimating the outcrossing rate of barley landraces and wild barley populations collected from ecologically different regions of Jordan. Theor Appl Genet 109:588–595PubMedCrossRefGoogle Scholar
  2. Aukerman MJ, Sakai H (2003) Regulation of flowering time and floral organ identity by a microRNA and its APETALA2-like target genes. Plant Cell 15:2730–2741PubMedCrossRefGoogle Scholar
  3. Briggs D (1978) Barley. Chapman and Hall, LondonCrossRefGoogle Scholar
  4. Ceccarelli S (1978) Single-gene inheritance of anther extrusion in barley. J Hered 69:210–211Google Scholar
  5. Cheignon M (1972) Structural modifications of cell-walls during elongation of stamen filament of Zea mays L. Cr Acad Sci D Nat 275:549Google Scholar
  6. Cheignon M, Schaever J, Cornier N (1973) Elongation of stamen filament of graminaceae as growth phenomenon. Cr Acad Sci D Nat 276:319–322Google Scholar
  7. Chen X (2004) A microRNA as a translational repressor of APETALA2 in Arabidopsis flower development. Science 303:2022–2025PubMedCrossRefGoogle Scholar
  8. Chen A, Brûlé-Babel A, Baumann U, Collins NC (2009) Structure–function analysis of the barley genome: the gene-rich region of chromosome 2HL. Funct Integr Genomics 9:67–79PubMedCrossRefGoogle Scholar
  9. Dahleen LS, Morgan W, Mittal S, Bregitzer P, Brown RH, Hill NS (2012) Quantitative trait loci (QTL) for Fusarium ELISA compared to QTL for Fusarium head blight resistance and deoxynivalenol content in barley. Plant Breed 131:237–243CrossRefGoogle Scholar
  10. Daniell H (2002) Molecular strategies for gene containment in transgenic crops. Nat Biotechnol 20:581–586PubMedGoogle Scholar
  11. Heslop-Harrison Y, Heslop-Harrison JS (1996) Lodicule function and filament extension in the grasses: potassium ion movement and tissue specialization. Ann Bot-Lond 77:573–582CrossRefGoogle Scholar
  12. Honda I, Turuspekov Y, Komatsuda T, Watanabe Y (2005) Morphological and physiological analysis of cleistogamy in barley (Hordeum vulgare). Physiol Plant 124:524–531CrossRefGoogle Scholar
  13. Hori K, Kobayashi T, Sato K, Takeda K (2005) QTL analysis of Fusarium head blight resistance using a high-density linkage map in barley. Theor Appl Genet 111:1661–1672PubMedCrossRefGoogle Scholar
  14. Hori K, Sato K, Kobayashi T, Takeda K (2006) QTL analysis of fusarium head blight severity in recombinant inbred population derived from a cross between two-rowed barley varieties. Breed Sci 56:25–30CrossRefGoogle Scholar
  15. Kirby EJM, Appleyard M (1981) Cereal development guide. Cereal Unit, KenilworthGoogle Scholar
  16. Koevenig JL (1973) Floral development and stamen filament elongation in Cleome hassleriana. Am J Bot 60:122–129CrossRefGoogle Scholar
  17. Komatsuda T, Nakamura I, Takaiwa F, Oka S (1998) Development of STS markers closely linked to the vrs1 locus in barley, Hordeum vulgare. Genome 41:680–685Google Scholar
  18. Kurauchi N, Makino T, Hirose S (1994) Inhieritance of cleistogamy-chasmogamy in barley. Barley Genet Newsl 23:19Google Scholar
  19. Lord EM (1981) Cleistogamy—a tool for the study of floral morphogenesis, function and evolution. Bot Rev 47:421–449CrossRefGoogle Scholar
  20. Lu Q, Lillemo M, Skinnes H, He X, Shi J, Ji F, Dong Y, Bjornstad A (2012) Anther extrusion and plant height are associated with Type I resistance to Fusarium head blight in bread wheat line ‘Shanghai-3/Catbird’. Theor Appl Genet 126:317–334PubMedCrossRefGoogle Scholar
  21. Ma SM, Wang YF (2004) Molecular strategies for decreasing the gene flow of transgenic plants. Yi Chuan 26:556–559PubMedGoogle Scholar
  22. Mano Y, Kawasaki S, Takaiwa F, Komatsuda T (2001) Construction of a genetic map of barley (Hordeum vulgare L.) cross ‘Azumamugi’ ×  ‘Kanto Nakate Gold’ using a simple and efficient amplified fragment-length polymorphism system. Genome 44:284–292PubMedGoogle Scholar
  23. Nair SK, Wang N, Turuspekov Y, Pourkheirandish M, Sinsuwongwat S, Chen GX, Sameri M, Tagiri A, Honda I, Watanabe Y, Kanamori H, Wicker T, Stein N, Nagamura Y, Matsumoto T, Komatsuda T (2010) Cleistogamous flowering in barley arises from the suppression of microRNA-guided HvAP2 mRNA cleavage. Proc Natl Acad Sci USA 107:490–495PubMedCrossRefGoogle Scholar
  24. Sameri M, Takeda K, Komatsuda T (2006) Quantitative trait loci controlling agronomic traits in recombinant inbred lines from a cross of oriental- and occidental-type barley cultivars. Breed Sci 56:243–252CrossRefGoogle Scholar
  25. Sameri M, Nakamura S, Nair SK, Takeda K, Komatsuda T (2009) A quantitative trait locus for reduced culm internode length in barley segregates as a Mendelian gene. Theor Appl Genet 118:643–652PubMedCrossRefGoogle Scholar
  26. Simons KJ, Fellers JP, Trick HN, Zhang ZC, Tai YS, Gill BS, Faris JD (2006) Molecular characterization of the major wheat domestication gene Q. Genetics 172:547–555PubMedCrossRefGoogle Scholar
  27. Skinnes H, Semagn K, Tarkegne Y, Maroy AG, Bjornstad A (2010) The inheritance of anther extrusion in hexaploid wheat and its relationship to Fusarium head blight resistance and deoxynivalenol content. Plant Breed 129:149–155CrossRefGoogle Scholar
  28. Turuspekov Y, Mano Y, Honda I, Kawada N, Watanabe Y, Komatsuda T (2004) Identification and mapping of cleistogamy genes in barley. Theor Appl Genet 109:480–487PubMedCrossRefGoogle Scholar
  29. Turuspekov Y, Kawada N, Honda I, Watanabe Y, Komatsuda T (2005) Identification and mapping of a QTL for rachis internode length associated with cleistogamy in barley. Plant Breed 124:542–545CrossRefGoogle Scholar
  30. Yoshida H (2012) Is the lodicule a petal: molecular evidence? Plant Sci 184:121–128PubMedCrossRefGoogle Scholar
  31. Yoshida H, Itoh J, Ohmori S, Miyoshi K, Horigome A, Uchida E, Kimizu M, Matsumura Y, Kusaba M, Satoh H, Nagato Y (2007) superwoman1-cleistogamy, a hopeful allele for gene containment in GM rice. Plant Biotechnol J 5:835–846PubMedCrossRefGoogle Scholar
  32. Zeng XC, Zhou X, Zhang W, Murofushi N, Kitahara T, Kamuro Y (1999) Opening of rice floret in rapid response to methyl jasmonate. J Plant Growth Regul 18:153–158PubMedCrossRefGoogle Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2013

Authors and Affiliations

  • Ning Wang
    • 1
  • Shunzong Ning
    • 1
    • 2
  • Mohammad Pourkheirandish
    • 1
  • Ichiro Honda
    • 3
  • Takao Komatsuda
    • 1
    Email author
  1. 1.National Institute of Agrobiological SciencesPlant Genome Research UnitTsukubaJapan
  2. 2.Graduate School of HorticultureChiba UniversityMatsudo, ChibaJapan
  3. 3.Department of BiotechnologyMaebashi Institute of TechnologyMaebashiJapan

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