Note to users. If you're seeing this message, it means that your browser cannot find this page's style/presentation instructions -- or possibly that you are using a browser that does not support current Web standards. Find out more about why this message is appearing, and what you can do to make your experience of our site the best it can be.

Science 12 March 2004:
Vol. 303. no. 5664, pp. 1640 - 1644
DOI: 10.1126/science.1094305
Prev | Table of Contents | Next

Research Articles

The Wheat VRN2 Gene Is a Flowering Repressor Down-Regulated by Vernalization

Liuling Yan,1* Artem Loukoianov,1 Ann Blechl,2 Gabriela Tranquilli,1{dagger} Wusirika Ramakrishna,3 Phillip SanMiguel,4 Jeffrey L. Bennetzen,5 Viviana Echenique,1{ddagger} Jorge Dubcovsky1*§

Plants with a winter growth habit flower earlier when exposed for several weeks to cold temperatures, a process called vernalization. We report here the positional cloning of the wheat vernalization gene VRN2, a dominant repressor of flowering that is down-regulated by vernalization. Loss of function of VRN2, whether by natural mutations or deletions, resulted in spring lines, which do not require vernalization to flower. Reduction of the RNA level of VRN2 by RNA interference accelerated the flowering time of transgenic winter-wheat plants by more than a month.

1 Department of Agronomy and Range Science, University of California, Davis, CA 95616, USA.
2 U.S. Department of Agriculture–Agricultural Research Service, Albany, CA 94710, USA.
3 Department of Biological Sciences, Michigan Technological University, Houghton, MI 49931, USA.
4 Purdue University Genomics Core, Purdue University, West Lafayette, IN 47907, USA.
5 Department of Genetics, University of Georgia, Athens, GA 30602, USA.



* These authors contributed equally to this work.

{dagger} Present address: Instituto Recursos Biológicos, Instituto Nacional de Tecnologia Agropecuaria, (1712) Castelar, Buenos Aires, Argentina.

{ddagger} Present address: Consejo Nacional Investigaciones Cientificas y Tecnicas Departamento de Agronomía, Universidad Nacional del Sur, 8000 Bahía Blanca, Argentina.

§ To whom correspondence should be addressed. E-mail: jdubcovsky{at}ucdavis.edu

Read the Full Text


THIS ARTICLE HAS BEEN CITED BY OTHER ARTICLES:
Cis-regulatory Changes at FLOWERING LOCUS T Mediate Natural Variation in Flowering Responses of Arabidopsis thaliana.
C. Schwartz, S. Balasubramanian, N. Warthmann, T. P. Michael, J. Lempe, S. Sureshkumar, Y. Kobayashi, J. N. Maloof, J. O. Borevitz, J. Chory, et al. (2009)
Genetics 183, 723-732
   Abstract »    Full Text »    PDF »
Gene Content and Virtual Gene Order of Barley Chromosome 1H.
K. F.X. Mayer, S. Taudien, M. Martis, H. Simkova, P. Suchankova, H. Gundlach, T. Wicker, A. Petzold, M. Felder, B. Steuernagel, et al. (2009)
Plant Physiology 151, 496-505
   Abstract »    Full Text »    PDF »
Genome comparisons reveal a dominant mechanism of chromosome number reduction in grasses and accelerated genome evolution in Triticeae.
M. C. Luo, K. R. Deal, E. D. Akhunov, A. R. Akhunova, O. D. Anderson, J. A. Anderson, N. Blake, M. T. Clegg, D. Coleman-Derr, E. J. Conley, et al. (2009)
PNAS 106, 15780-15785
   Abstract »    Full Text »    PDF »
Identification of genomic regions determining the phenological development leading to floral transition in wheat (Triticum aestivum L.).
M. Baga, D. B. Fowler, and R. N. Chibbar (2009)
J. Exp. Bot. 60, 3575-3585
   Abstract »    Full Text »    PDF »
Molecular Evolution of VEF-Domain-Containing PcG Genes in Plants.
L.-J. Chen, Z.-Y. Diao, C. Specht, and Z. R. Sung (2009)
Mol Plant 2, 738-754
   Abstract »    Full Text »    PDF »
What Has Natural Variation Taught Us about Plant Development, Physiology, and Adaptation?.
C. Alonso-Blanco, M. G.M. Aarts, L. Bentsink, J. J.B. Keurentjes, M. Reymond, D. Vreugdenhil, and M. Koornneef (2009)
PLANT CELL 21, 1877-1896
   Abstract »    Full Text »    PDF »
The molecular biology of seasonal flowering-responses in Arabidopsis and the cereals.
A. Greenup, W. J. Peacock, E. S. Dennis, and B. Trevaskis (2009)
Ann. Bot. 103, 1165-1172
   Abstract »    Full Text »    PDF »
Vernalization-induced flowering in cereals is associated with changes in histone methylation at the VERNALIZATION1 gene.
S. N. Oliver, E. J. Finnegan, E. S. Dennis, W. J. Peacock, and B. Trevaskis (2009)
PNAS 106, 8386-8391
   Abstract »    Full Text »    PDF »
The CArG-Box Located Upstream from the Transcriptional Start of Wheat Vernalization Gene VRN1 Is Not Necessary for the Vernalization Response.
B. Pidal, L. Yan, D. Fu, F. Zhang, G. Tranquilli, and J. Dubcovsky (2009)
J. Hered. 100, 355-364
   Abstract »    Full Text »    PDF »
The influence of vernalization and daylength on expression of flowering-time genes in the shoot apex and leaves of barley (Hordeum vulgare)..
S. Sasani, M. N. Hemming, S. N. Oliver, A. Greenup, R. Tavakkol-Afshari, S. Mahfoozi, K. Poustini, H.-R. Sharifi, E. S. Dennis, W. J. Peacock, et al. (2009)
J. Exp. Bot. 60, 2169-2178
   Abstract »    Full Text »    PDF »
Development of 5006 Full-Length CDNAs in Barley: A Tool for Accessing Cereal Genomics Resources.
K. Sato, T. Shin-I, M. Seki, K. Shinozaki, H. Yoshida, K. Takeda, Y. Yamazaki, M. Conte, and Y. Kohara (2009)
DNA Res 16, 81-89
   Abstract »    Full Text »    PDF »
PCR-Based Markers Diagnostic for Spring and Winter Seasonal Growth Habit in Barley.
J. Cockram, C. Norris, and D. M. O'Sullivan (2009)
Crop Sci. 49, 403-410
   Abstract »    Full Text »    PDF »
Genetic and Molecular Characterization of the VRN2 Loci in Tetraploid Wheat.
A. Distelfeld, G. Tranquilli, C. Li, L. Yan, and J. Dubcovsky (2009)
Plant Physiology 149, 245-257
   Abstract »    Full Text »    PDF »
Independent Losses of Function in a Polyphenol Oxidase in Rice: Differentiation in Grain Discoloration between Subspecies and the Role of Positive Selection under Domestication.
Y. Yu, T. Tang, Q. Qian, Y. Wang, M. Yan, D. Zeng, B. Han, C.-I Wu, S. Shi, and J. Li (2008)
PLANT CELL 20, 2946-2959
   Abstract »    Full Text »    PDF »
The chromosome region including the earliness per se locus Eps-Am1 affects the duration of early developmental phases and spikelet number in diploid wheat.
S. Lewis, M. E. Faricelli, M. L. Appendino, M. Valarik, and J. Dubcovsky (2008)
J. Exp. Bot. 59, 3595-3607
   Abstract »    Full Text »    PDF »
Regulation of floral initiation in horticultural trees.
J. D. Wilkie, M. Sedgley, and T. Olesen (2008)
J. Exp. Bot. 59, 3215-3228
   Abstract »    Full Text »    PDF »
Involvement of the MADS-Box Gene ZMM4 in Floral Induction and Inflorescence Development in Maize.
O. N. Danilevskaya, X. Meng, D. A. Selinger, S. Deschamps, P. Hermon, G. Vansant, R. Gupta, E. V. Ananiev, and M. G. Muszynski (2008)
Plant Physiology 147, 2054-2069
   Abstract »    Full Text »    PDF »
Effects of photo and thermo cycles on flowering time in barley: a genetical phenomics approach.
I. Karsai, P. Szucs, B. Koszegi, P.M. Hayes, A. Casas, Z. Bedo, and O. Veisz (2008)
J. Exp. Bot. 59, 2707-2715
   Abstract »    Full Text »    PDF »
Molecular Plant Breeding as the Foundation for 21st Century Crop Improvement.
S. P. Moose and R. H. Mumm (2008)
Plant Physiology 147, 969-977
   Full Text »    PDF »
Low-Temperature and Daylength Cues Are Integrated to Regulate FLOWERING LOCUS T in Barley.
M. N. Hemming, W. J. Peacock, E. S. Dennis, and B. Trevaskis (2008)
Plant Physiology 147, 355-366
   Abstract »    Full Text »    PDF »
FLC or not FLC: the other side of vernalization.
C. M. Alexandre and L. Hennig (2008)
J. Exp. Bot.
   Abstract »    Full Text »    PDF »
Natural Variation in Arabidopsis lyrata Vernalization Requirement Conferred by a FRIGIDA Indel Polymorphism.
H. Kuittinen, A. Niittyvuopio, P. Rinne, and O. Savolainen (2008)
Mol. Biol. Evol. 25, 319-329
   Abstract »    Full Text »    PDF »
Discrete Developmental Roles for Temperate Cereal Grass VERNALIZATION1/FRUITFULL-Like Genes in Flowering Competency and the Transition to Flowering.
J. C. Preston and E. A. Kellogg (2008)
Plant Physiology 146, 265-276
   Abstract »    Full Text »    PDF »
The molecular analysis of the shade avoidance syndrome in the grasses has begun.
T. H. Kebrom and T. P. Brutnell (2007)
J. Exp. Bot.
   Abstract »    Full Text »    PDF »
The Importance of Barley Genetics and Domestication in a Global Perspective.
M. Pourkheirandish and T. Komatsuda (2007)
Ann. Bot. 100, 999-1008
   Abstract »    Full Text »    PDF »
Overexpression of TaVRN1 in Arabidopsis Promotes Early Flowering and Alters Development.
H. Adam, F. Ouellet, N. A. Kane, Z. Agharbaoui, G. Major, Y. Tominaga, and F. Sarhan (2007)
Plant Cell Physiol. 48, 1192-1206
   Abstract »    Full Text »    PDF »
Genome Plasticity a Key Factor in the Success of Polyploid Wheat Under Domestication.
J. Dubcovsky and J. Dvorak (2007)
Science 316, 1862-1866
   Abstract »    Full Text »    PDF »
Evolutionary Conservation of the FLOWERING LOCUS C-Mediated Vernalization Response: Evidence From the Sugar Beet (Beta vulgaris).
P. A. Reeves, Y. He, R. J. Schmitz, R. M. Amasino, L. W. Panella, and C. M. Richards (2007)
Genetics 176, 295-307
   Abstract »    Full Text »    PDF »
Control of flowering time in temperate cereals: genes, domestication, and sustainable productivity.
J. Cockram, H. Jones, F. J. Leigh, D. O'Sullivan, W. Powell, D. A. Laurie, and A. J. Greenland (2007)
J. Exp. Bot.
   Abstract »    Full Text »    PDF »
Short Vegetative Phase-Like MADS-Box Genes Inhibit Floral Meristem Identity in Barley.
B. Trevaskis, M. Tadege, M. N. Hemming, W. J. Peacock, E. S. Dennis, and C. Sheldon (2007)
Plant Physiology 143, 225-235
   Abstract »    Full Text »    PDF »
From the Cover: The wheat and barley vernalization gene VRN3 is an orthologue of FT.
L. Yan, D. Fu, C. Li, A. Blechl, G. Tranquilli, M. Bonafede, A. Sanchez, M. Valarik, S. Yasuda, and J. Dubcovsky (2006)
PNAS 103, 19581-19586
   Abstract »    Full Text »    PDF »
delayed flowering1 Encodes a Basic Leucine Zipper Protein That Mediates Floral Inductive Signals at the Shoot Apex in Maize.
M. G. Muszynski, T. Dam, B. Li, D. M. Shirbroun, Z. Hou, E. Bruggemann, R. Archibald, E. V. Ananiev, and O. N. Danilevskaya (2006)
Plant Physiology 142, 1523-1536
   Abstract »    Full Text »    PDF »
Comparative Mapping of Growth Habit, Plant Height, and Flowering QTLs in Two Interspecific Families of Leymus.
S. R. Larson, X. Wu, T. A. Jones, K. B. Jensen, N. J. Chatterton, B. L. Waldron, J. G. Robins, B. S. Bushman, and A. J. Palazzo (2006)
Crop Sci. 46, 2526-2539
   Abstract »    Full Text »    PDF »
CONSTANS and the CCAAT Box Binding Complex Share a Functionally Important Domain and Interact to Regulate Flowering of Arabidopsis.
S. Wenkel, F. Turck, K. Singer, L. Gissot, J. Le Gourrierec, A. Samach, and G. Coupland (2006)
PLANT CELL 18, 2971-2984
   Abstract »    Full Text »    PDF »
RNA Interference-Based Gene Silencing as an Efficient Tool for Functional Genomics in Hexaploid Bread Wheat.
S. Travella, T. E. Klimm, and B. Keller (2006)
Plant Physiology 142, 6-20
   Abstract »    Full Text »    PDF »
High-Resolution Radiation Hybrid Map of Wheat Chromosome 1D.
V. Kalavacharla, K. Hossain, Y. Gu, O. Riera-Lizarazu, M. I. Vales, S. Bhamidimarri, J. L. Gonzalez-Hernandez, S. S. Maan, and S. F. Kianian (2006)
Genetics 173, 1089-1099
   Abstract »    Full Text »    PDF »
The transcription factor FLC confers a flowering response to vernalization by repressing meristem competence and systemic signaling in Arabidopsis..
I. Searle, Y. He, F. Turck, C. Vincent, F. Fornara, S. Krober, R. A. Amasino, and G. Coupland (2006)
Genes & Dev. 20, 898-912
   Abstract »    Full Text »    PDF »
Characterization of the Vernalization Response in Lolium perenne by a cDNA Microarray Approach.
S. Ciannamea, J. Busscher-Lange, S. de Folter, G. C. Angenent, and R. G. H. Immink (2006)
Plant Cell Physiol. 47, 481-492
   Abstract »    Full Text »    PDF »
HvVRN2 Responds to Daylength, whereas HvVRN1 Is Regulated by Vernalization and Developmental Status.
B. Trevaskis, M. N. Hemming, W. J. Peacock, and E. S. Dennis (2006)
Plant Physiology 140, 1397-1405
   Abstract »    Full Text »    PDF »
TaVRT-2, a Member of the StMADS-11 Clade of Flowering Repressors, Is Regulated by Vernalization and Photoperiod in Wheat.
N. A. Kane, J. Danyluk, G. Tardif, F. Ouellet, J.-F. Laliberte, A. E. Limin, D. B. Fowler, and F. Sarhan (2005)
Plant Physiology 138, 2354-2363
   Abstract »    Full Text »    PDF »
Regulation of VRN-1 Vernalization Genes in Normal and Transgenic Polyploid Wheat.
A. Loukoianov, L. Yan, A. Blechl, A. Sanchez, and J. Dubcovsky (2005)
Plant Physiology 138, 2364-2373
   Abstract »    Full Text »    PDF »
DNA Rearrangement in Orthologous Orp Regions of the Maize, Rice and Sorghum Genomes.
J. Ma, P. SanMiguel, J. Lai, J. Messing, and J. L. Bennetzen (2005)
Genetics 170, 1209-1220
   Abstract »    Full Text »    PDF »
Flowering of the Grass Lolium perenne. Effects of Vernalization and Long Days on Gibberellin Biosynthesis and Signaling.
C. P. MacMillan, C. A. Blundell, and R. W. King (2005)
Plant Physiology 138, 1794-1806
   Abstract »    Full Text »    PDF »
Conservation of Arabidopsis Flowering Genes in Model Legumes.
V. Hecht, F. Foucher, C. Ferrandiz, R. Macknight, C. Navarro, J. Morin, M. E. Vardy, N. Ellis, J. P. Beltran, C. Rameau, et al. (2005)
Plant Physiology 137, 1420-1434
   Abstract »    Full Text »    PDF »
Abiotic Stress Tolerance in Grasses. From Model Plants to Crop Plants.
M. Tester and A. Bacic (2005)
Plant Physiology 137, 791-793
   Full Text »    PDF »
Low levels of linkage disequilibrium in wild barley (Hordeum vulgare ssp. spontaneum) despite high rates of self-fertilization.
P. L. Morrell, D. M. Toleno, K. E. Lundy, and M. T. Clegg (2005)
PNAS 102, 2442-2447
   Abstract »    Full Text »    PDF »
Fine Mapping of a Grain-Weight Quantitative Trait Locus in the Pericentromeric Region of Rice Chromosome 3.
J. Li, M. Thomson, and S. R. McCouch (2004)
Genetics 168, 2187-2195
   Abstract »    Full Text »    PDF »
Marker-Assisted Selection in Public Breeding Programs: The Wheat Experience.
J. Dubcovsky (2004)
Crop Sci. 44, 1895-1898
   Full Text »    PDF »
A 2500-Locus Bin Map of Wheat Homoeologous Group 5 Provides Insights on Gene Distribution and Colinearity With Rice.
A. M. Linkiewicz, L. L. Qi, B. S. Gill, A. Ratnasiri, B. Echalier, S. Chao, G. R. Lazo, D. D. Hummel, O. D. Anderson, E. D. Akhunov, et al. (2004)
Genetics 168, 665-676
   Abstract »    Full Text »    PDF »
A Workshop Report on Wheat Genome Sequencing: International Genome Research on Wheat Consortium.
B. S. Gill, R. Appels, A.-M. Botha-Oberholster, C. R. Buell, J. L. Bennetzen, B. Chalhoub, F. Chumley, J. Dvorak, M. Iwanaga, B. Keller, et al. (2004)
Genetics 168, 1087-1096
   Abstract »    Full Text »    PDF »
Vernalization, Competence, and the Epigenetic Memory of Winter.
R. Amasino (2004)
PLANT CELL 16, 2553-2559
   Full Text »    PDF »
Control of Arabidopsis flowering: the chill before the bloom.
I. R. Henderson and C. Dean (2004)
Development 131, 3829-3838
   Abstract »    Full Text »    PDF »
Positioning Arabidopsis in Plant Biology. A Key Step Toward Unification of Plant Research.
M. Bevan and S. Walsh (2004)
Plant Physiology 135, 602-606
   Abstract »    Full Text »    PDF »


To Advertise     Find Products

Science. ISSN 0036-8075 (print), 1095-9203 (online)