Abstract
With the aim of dissecting the genetic determinants of flowering time, vernalization response, and photoperiod sensitivity, we mapped the candidate genes for Vrn-H2 and Vrn-H1 in a facultative × winter barley mapping population and determined their relationships with flowering time and vernalization via QTL analysis. The Vrn-H2 candidate ZCCT-H genes were completely missing from the facultative parent and present in the winter barley parent. This gene was the major determinant of flowering time under long photoperiods in controlled environment experiments, irrespective of vernalization, and under spring-sown field experiments. It was the sole determinant of vernalization response, but the effect of the deletion was modulated by photoperiods when the vernalization requirement was fulfilled. There was no effect under short photoperiods. The Vrn-H1 candidate gene (HvBM5A) was mapped based on a microsatellite polymorphism we identified in the promoter of this gene. Otherwise, the HvBM5A alleles for the two parents were identical. Therefore, the significant flowering time QTL effect associated with this locus suggests tight linkage rather than pleiotropy. This QTL effect was smaller in magnitude than those associated with the Vrn-H2 locus and was significant in two-way interactions with Vrn-H2. The Vrn-H1 locus had no effect on vernalization response. Our results support the Vrn-H2/Vrn-H1 repressor/structural gene model for vernalization response in barley and suggest that photoperiod may also affect the Vrn genes or tightly linked loci.
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Danyluk J, Kane ND, Breton G, Limin AE, Fowler DB, Sarhan F (2003) TaVRT-1, a putative transcription factor associated with vegetative to reproductive transition in cereals. Plant Physiol 132:1849–1860
Dubcovsky J, Lijavetzky D, Appendino L, Tranquilli G (1998) Comparative RFLP mapping of Triticum monococcum genes controlling vernalization requirement. Theor Appl Genet 97:968–975
Dubcovsky J, Chen C, Yan L (2005) Molecular characterization of the allelic variation at the VRN-H2 vernalization locus in barley. Mol Breed (in press)
Fankhauser C, Staiger D (2002) Photoreceptors in Arabidopsis thaliana: light perception, signal transduction and entrainment of the endogenous clock. Planta 216:1–16
Fu D, Szűcs P, Yan L, Helguera M, Skinner JS, von Zitzewitz J, Hayes PM, Dubcovsky J (2005) Large deletions within the VRN-1 first intron are associated with spring growth habit in barley and wheat. Mol Genet Genomics (in press) DOI: 10.1007/s00438-004-1095-4.
Galiba G, Quarrie SA, Sutka J, Morgounov A, Snape JW (1995) RFLP mapping of vernalization (Vrn1) and frost resistance (Fr1) genes on chromosome 5A of wheat. Theor Appl Genet 90:1174–1179
Hackett CA, Ellis RP, Forster BP, McNicol JW, Macaulay M (1992) Statistical analysis of a linkage experiment in barley involving quantitative trait loci for height and ear-emergence time and two genetic markers on chromosome 4. Theor Appl Genet 85:120–126
Hayes PM, Liu BH, Knapp SJ, Chen F, Jones B, Blake T, Frankowiak JD, Rasmusson D, Sorrells ME, Ullrich SE, Wesenberg D, Kleinhofs A (1993) Quantitative trait locus effects and environmental interaction in a sample of North American barley germplasm. Theor Appl Genet 87:392–401
Karsai I, Mészáros K, Hayes PM, Bedő Z (1997) Effects of loci on chromosomes 2 (2H) and 7 (5H) on developmental patterns in barley (Hordeum vulgare L.) under different photoperiod regimes. Theor Appl Genet 94:612–618
Karsai I, Mészáros K, Láng L, Hayes PM, Bedő Z (2001) Multivariate analysis of traits determining adaptation in cultivated barley. Plant Breed 120:217–222
Karsai I, Mészáros K, Szűcs P, Filichkina T, Hayes PM, Láng L, Bedő Z (2004a) Effect of Vrn-H2 vernalization response locus (4H) on plant development in a winter barley mapping population. In: Proceedings of the 9th barley genetic conference, June 20–26, Brno, pp 289–294
Karsai I, Hayes PM, Kling J, Matus IA, Mészáros K, Láng L, Bedő Z, Sato K (2004b) Genetic variation in component traits of flowering time in Hordeum vulgare subsp. spontaneum accessions characterized in controlled environments. Crop Sci 44:1622–1632
Kato K, Miura H, Sawada S (1999) QTL mapping of genes controlling ear emergence time and plant height on chromosome 5A of wheat. Theor Appl Genet 98:472–477
Lander ES, Botstein D (1989) Mapping Mendelian factors underlying quantitative traits using RFLP linkage maps. Genetics 121:185–199
Lander ES, Green P, Abrahamson J, Barlow A, Daly MJ, Lincoln SE, Newburg L (1987) MAPMAKER: an interaction computer package for constructing primary genetic linkage maps for experimental and natural populations. Genomics 1:174–181
Laurie DA, Pratchett N, Bezant JH, Snape JW (1994) Genetic analysis of a photoperiod response gene on the short arm of chromosome 2(2H) of Hordeum vulgare (barley). Heredity 72:619–627
Laurie DA, Pratchett N, Bezant JH, Snape JW (1995) RFLP mapping of five major genes and eight quantitative trait loci controlling flowering time in a winter × spring barley (Hordeum vulgare L.) cross. Genome 38:575–585
Law CN, Worland AJ, Giorgi B (1976) The genetic control of ear-emergence time by chromosomes 5A and 5D of wheat. Heredity 36:49–58
Pan A, Hayes PM, Chen F, Chen THH, Blake T, Wright S, Karsai I, Bedő Z (1994) Genetic analysis of the components of winterhardiness in barley (Hordeum vulgare L.). Theor Appl Genet 89:900–910
Pillen K, Binder A, Kreuzkam B, Ramsay L, Waugh R, Förster J, Léon J (2000) Mapping of EMBL-derived barley microsatellites and their use to differentiate German barley cultivars. Theor Appl Genet 101:652–660
Ramsay L, Macaulay M, degli Ivanissevich S, MacLean K, Cardle L, Fuller J, Edwards KJ, Tuvesson S, Morgante M, Massari A, Maestri E, Marmiroli N, Sjakste T, Ganal M, Powell W, Waugh R (2000) A simple sequence repeat-based linkage map of barley. Genetics 156:1997–2005
Takahashi R, Yasuda S (1971) Genetics of earliness and growth habit in barley. In: Nilan RA (ed) Barley genetics II. Washington State University Press, Pullman, pp 388–408
Tottman DR, Makepeace RJ (1979) An explanation of the decimal code for the growth stages of cereals, with illustrations. Ann Appl Biol 93:221–234
Tranquilli G, Dubcovsky J (2000) Epistatic interaction between vernalization genes Vrn-Am 1 and Vrn-Am 2 in diploid wheat. J Hered 91:304–306
Yan L, Echenique V, Busso C, SanMiguel P, Ramakrishna W, Bennetzen JL, Harrington S, Dubcovsky J (2002) Cereal genes similar to Snf2 define a new subfamily that includes human and mouse genes. Mol Genet Genomics 268:488–499
Yan L, Loukoianov A, Tranquilli G, Helguera M, Fahima T, Dubcovsky J (2003) Positional cloning of the wheat vernalization gene VRN1. PNAS 100:6263–6268
Yan L, Loukoianov A, Blechl A, Tranquilli G, Ramakrishna W, SanMiguel P, Bennetzen JL, Echenique V, Dubcovsky J (2004) The wheat VRN2 gene is a flowering repressor down-regulated by vernalization. Science 303:1640–1644
Yasuda S, Hayashi J, Moriya I (1993) Genetic constitution for spring growth habit and some other characters in barley cultivars in the Mediterranean coastal regions. Euphytica 70:77–83
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This project was funded by the Hungarian National Research and Innovation Fund (NKFP) research grant no. 4/0020/2002 and the United States National Science Foundation project no. DBIO 110124.
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Communicated by J.W. Snape
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Karsai, I., Szűcs, P., Mészáros, K. et al. The Vrn-H2 locus is a major determinant of flowering time in a facultative × winter growth habit barley (Hordeum vulgare L.) mapping population. Theor Appl Genet 110, 1458–1466 (2005). https://doi.org/10.1007/s00122-005-1979-7
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DOI: https://doi.org/10.1007/s00122-005-1979-7