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Richard M Amasino - One of the best experts on this subject based on the ideXlab platform.
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an ortholog of curly leaf enhancer of zeste like 1 is required for proper flowering in brachypodium distachyon
Plant Journal, 2018Co-Authors: Aaron Lomax, Daniel P Woods, Yinxin Dong, Frederic Bouche, Ying Rong, Kevin S Mayer, Xuehua Zhong, Richard M AmasinoAbstract:Many plants require prolonged exposure to cold to acquire the competence to flower. The process by which cold exposure results in competence is known as vernalization. In Arabidopsis thaliana, vernalization leads to the stable repression of the floral repressor FLOWERING LOCUS C via chromatin modification, including an increase of trimethylation on lysine 27 of histone H3 (H3K27me3) by Polycomb Repressive Complex 2 (PRC2). Vernalization in pooids is associated with the stable induction of a floral promoter, VERNALIZATION 1 (VRN1). From a screen for mutants with a reduced vernalization requirement in the model grass Brachypodium distachyon, we identified two recessive alleles of ENHANCER OF ZESTE-LIKE 1 (EZL1). EZL1 is orthologous to A. thaliana CURLY LEAF 1, a gene that encodes the catalytic subunit of PRC2. B. distachyon ezl1 mutants flower rapidly without vernalization in long-day (LD) photoperiods; thus, EZL1 is required for the proper maintenance of the vegetative state prior to vernalization. Transcriptomic studies in ezl1 revealed mis-regulation of thousands of genes, including ectopic expression of several floral homeotic genes in leaves. Loss of EZL1 results in the global reduction of H3K27me3 and H3K27me2, consistent with this gene making a major contribution to PRC2 activity in B. distachyon. Furthermore, in ezl1 mutants, the flowering genes VRN1 and AGAMOUS (AG) are ectopically expressed and have reduced H3K27me3. Artificial microRNA knock-down of either VRN1 or AG in ezl1-1 mutants partially restores wild-type flowering behavior in non-vernalized plants, suggesting that ectopic expression in ezl1 mutants may contribute to the rapid-flowering phenotype.
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a methyltransferase required for proper timing of the vernalization response in arabidopsis
Proceedings of the National Academy of Sciences of the United States of America, 2015Co-Authors: Wei Zhao, Wenhui Shen, Richard M AmasinoAbstract:Prolonged exposure to winter cold enables flowering in many plant species through a process called vernalization. In Arabidopsis, vernalization results from the epigenetic silencing of the floral repressor FLOWERING LOCUS C (FLC) via a Polycomb Repressive Complex 2 (PRC2)-mediated increase in the density of the epigenetic silencing mark H3K27me3 at FLC chromatin. During cold exposure, a gene encoding a unique, cold-specific PRC2 component, VERNALIZATION INSENSITIVE 3 (VIN3), which is necessary for PRC2-mediated silencing of FLC, is induced. Here we show that SET DOMAIN GROUP 7 (SDG7) is required for proper timing of VIN3 induction and of the vernalization process. Loss of SDG7 results in a vernalization-hypersensitive phenotype, as well as more rapid cold-mediated up-regulation of VIN3. In the absence of cold, loss of SDG7 results in elevated levels of long noncoding RNAs, which are thought to participate in epigenetic repression of FLC. Furthermore, loss of SDG7 results in increased H3K27me3 deposition on FLC chromatin in the absence of cold exposure and enhanced H3K27me3 spreading during cold treatment. Thus, SDG7 is a negative regulator of vernalization, and loss of SDG7 creates a partially vernalized state without cold exposure.
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the molecular basis of vernalization in different plant groups
Cold Spring Harbor Symposia on Quantitative Biology, 2012Co-Authors: Thomas S Ream, Daniel P Woods, Richard M AmasinoAbstract:: Timing of flowering is key to the reproductive success of many plants. In temperate climates, flowering is often coordinated with seasonal environmental cues such as temperature and photoperiod. Vernalization, the process by which a prolonged exposure to the cold of winter results in competence to flower during the following spring, is an example of the influence of temperature on the timing of flowering. In different groups of plants, there are distinct genes involved in vernalization, indicating that vernalization systems evolved independently in different plant groups. The convergent evolution of vernalization systems is not surprising given that angiosperm families had begun to diverge in warmer paleoclimates in which a vernalization response was not advantageous. Here, we review what is known of the vernalization response in three different plant groups: crucifers (Arabidopsis), Amaranthaceae (sugar beet), and Pooideae (wheat, barley, and Brachypodium distachyon). We also discuss the advantages of using Brachypodium as a model system to study flowering and vernalization in the Pooids. Finally, we discuss the evolution and function of the Ghd7/VRN2 gene family in grasses.
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vernalization winter and the timing of flowering in plants
Annual Review of Cell and Developmental Biology, 2009Co-Authors: Mark R Doyle, Sibum Sung, Richard M AmasinoAbstract:Plants have evolved many systems to sense their environment and to modify their growth and development accordingly. One example is vernalization, the process by which flowering is promoted as plants sense exposure to the cold temperatures of winter. A requirement for vernalization is an adaptive trait that helps prevent flowering before winter and permits flowering in the favorable conditions of spring. In Arabidopsis and cereals, vernalization results in the suppression of genes that repress flowering. We describe recent progress in understanding the molecular basis of this suppression. In Arabidopsis, vernalization involves the recruitment of chromatin-modifying complexes to a clade of flowering repressors that are silenced epigenetically via histone modifications. We also discuss the similarities and differences in vernalization between Arabidopsis and cereals.
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histone arginine methylation is required for vernalization induced epigenetic silencing of flc in winter annual arabidopsis thaliana
Proceedings of the National Academy of Sciences of the United States of America, 2008Co-Authors: Robert J Schmitz, Sibum Sung, Richard M AmasinoAbstract:Certain plant varieties typically require prolonged exposure to the cold of winter to become competent to flower rapidly in the spring. This process is known as vernalization. In Arabidopsis thaliana, vernalization renders plants competent to flower by epigenetically silencing the strong floral repressor FLOWERING LOCUS C (FLC). As a result of vernalization, levels of lysine-9 and lysine-27 trimethylation on histone 3, modifications that are characteristic of facultative heterochromatin in plants, increase at FLC chromatin. We have identified a mutant, protein arginine methyltransferase 5 (atprmt5), that fails to flower rapidly after vernalization treatment. AtPRMT5 encodes a type II protein arginine methyltransferase (PRMT) that, in winter-annual strains, is required for epigenetic silencing of FLC and for the vernalization-mediated histone modifications characteristic of the vernalized state. Furthermore, the levels of arginine methylation of FLC chromatin increase after vernalization. Therefore, arginine methylation of FLC chromatin is part of the histone code that is required for mitotic stability of the vernalized state.
Elizabeth S Dennis - One of the best experts on this subject based on the ideXlab platform.
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vernalization induced flowering in cereals is associated with changes in histone methylation at the vernalization1 gene
Proceedings of the National Academy of Sciences of the United States of America, 2009Co-Authors: Sandra N Oliver, E J Finnegan, W J Peacock, Elizabeth S Dennis, Ben TrevaskisAbstract:Prolonged exposure to low temperatures (vernalization) accelerates the transition to reproductive growth in many plant species, including the model plant Arabidopsis thaliana and the economically important cereal crops, wheat and barley. Vernalization-induced flowering is an epigenetic phenomenon. In Arabidopsis, stable down-regulation of FLOWERING LOCUS C (FLC) by vernalization is associated with changes in histone modifications at FLC chromatin. In cereals, the vernalization response is mediated by stable induction of the floral promoter VERNALIZATION1 (VRN1), which initiates reproductive development at the shoot apex. We show that in barley (Hordeum vulgare), repression of HvVRN1 before vernalization is associated with high levels of histone 3 lysine 27 trimethylation (H3K27me3) at HvVRN1 chromatin. Vernalization caused increased levels of histone 3 lysine 4 trimethylation (H3K4me3) and a loss of H3K27me3 at HvVRN1, suggesting that vernalization promotes an active chromatin state at VRN1. Levels of these histone modifications at 2 other flowering-time genes, VERNALIZATION2 and FLOWERING LOCUS T, were not altered by vernalization. Our study suggests that maintenance of an active chromatin state at VRN1 is likely to be the basis for epigenetic memory of vernalization in cereals. Thus, regulation of chromatin state is a feature of epigenetic memory of vernalization in Arabidopsis and the cereals; however, whereas vernalization-induced flowering in Arabidopsis is mediated by epigenetic regulation of the floral repressor FLC, this phenomenon in cereals is mediated by epigenetic regulation of the floral activator, VRN1.
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the influence of vernalization and daylength on expression of flowering time genes in the shoot apex and leaves of barley hordeum vulgare
Journal of Experimental Botany, 2009Co-Authors: Sandra N Oliver, Shahryar Sasani, Megan N Hemming, Aaron Greenup, Reza Tavakkolafshari, Siroos Mahfoozi, Hamidreza Sharifi, K Poustini, Elizabeth S DennisAbstract:Responses to prolonged low-temperature treatment of imbibed seeds (vernalization) were examined in barley (Hordeum vulgare). These occurred in two phases: the perception of prolonged cold, which occurred gradually at low temperatures, and the acceleration of reproductive development, which occurred after vernalization. Expression of the VERNALIZATION1 gene (HvVRN1) increased gradually in germinating seedlings during vernalization, both at the shoot apex and in the developing leaves. This occurred in darkness, independently of VERNALIZATION2 (HvVRN2), consistent with the hypothesis that expression of HvVRN1 is induced by prolonged cold independently of daylength flowering-response pathways. After vernalization, expression of HvVRN1 was maintained in the shoot apex and leaves. This was associated with accelerated inflorescence initiation and with down-regulation of HvVRN2 in the leaves. The largest determinant of HvVRN1 expression levels in vernalized plants was the length of seed vernalization treatment. Daylength did not influence HvVRN1 expression levels in shoot apices and typically did not affect expression in leaves. In the leaves of plants that had experienced a saturating seed vernalization treatment, expression of HvVRN1 was higher in long days, however. HvFT1 was expressed in the leaves of these plants in long days, which might account for the elevated HvVRN1 expression. Long-day up-regulation of HvVRN1 was not required for inflorescence initiation, but might accelerate subsequent stages of inflorescence development. Similar responses to seed vernalization were also observed in wheat (Triticum aestivum). These data support the hypothesis that VRN1 is induced by cold during winter to promote spring flowering in vernalization-responsive cereals.
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the molecular basis of vernalization induced flowering in cereals
Trends in Plant Science, 2007Co-Authors: Ben Trevaskis, Megan N Hemming, Elizabeth S Dennis, James W PeacockAbstract:Genetic analyses have identified three genes that control the vernalization requirement in wheat and barley; VRN1, VRN2 and FT (VRN3). These genes have now been isolated and shown to regulate not only the vernalization response but also the promotion of flowering by long days. VRN1 is induced by vernalization and accelerates the transition to reproductive development at the shoot apex. FT is induced by long days and further accelerates reproductive apex development. VRN2, a floral repressor, integrates vernalization and day-length responses by repressing FT until plants are vernalized. A comparison of flowering time pathways in cereals and Arabidopsis shows that the vernalization response is controlled by different MADS box genes, but integration of vernalization and long-day responses occurs through similar mechanisms.
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the arabidopsis thaliana vernalization response requires a polycomb like protein complex that also includes vernalization insensitive 3
Proceedings of the National Academy of Sciences of the United States of America, 2006Co-Authors: Craig C Wood, James W Peacock, Masumi Robertson, Greg Tanner, Elizabeth S Dennis, Chris A HelliwellAbstract:In Arabidopsis thaliana, the promotion of flowering by cold temperatures, vernalization, is regulated via a floral-repressive MADS box transcription factor, FLOWERING LOCUS C (FLC). Vernalization leads to the epigenetic repression of FLC expression, a process that requires the polycomb group (PcG) protein VERNALIZATION 2 (VRN2) and the plant homeodomain protein VERNALIZATION INSENSITIVE 3 (VIN3). We demonstrate that the repression of FLC by vernalization requires homologues of other Polycomb Repressive Complex 2 proteins and VRN2. We show in planta that VRN2 and VIN3 are part of a large protein complex that can include the PcG proteins FERTILIZATION INDEPENDENT ENDOSPERM, CURLY LEAF, and SWINGER. These findings suggest a single protein complex is responsible for histone deacetylation at FLC and histone methylation at FLC in vernalized plants. The abundance of the complex increases during vernalization and declines after plants are returned to higher temperatures, consistent with the complex having a role in establishing FLC repression.
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the arabidopsis thaliana vernalization response requires a polycomb like protein complex that also includes vernalization insensitive 3
Proceedings of the National Academy of Sciences of the United States of America, 2006Co-Authors: Craig C Wood, James W Peacock, Masumi Robertson, Greg Tanner, Elizabeth S Dennis, Chris A HelliwellAbstract:In Arabidopsis thaliana, the promotion of flowering by cold temperatures, vernalization, is regulated via a floral-repressive MADS box transcription factor, FLOWERING LOCUS C (FLC). Vernalization leads to the epigenetic repression of FLC expression, a process that requires the polycomb group (PcG) protein VERNALIZATION 2 (VRN2) and the plant homeodomain protein VERNALIZATION INSENSITIVE 3 (VIN3). We demonstrate that the repression of FLC by vernalization requires homologues of other Polycomb Repressive Complex 2 proteins and VRN2. We show in planta that VRN2 and VIN3 are part of a large protein complex that can include the PcG proteins FERTILIZATION INDEPENDENT ENDOSPERM, CURLY LEAF, and SWINGER. These findings suggest a single protein complex is responsible for histone deacetylation at FLC and histone methylation at FLC in vernalized plants. The abundance of the complex increases during vernalization and declines after plants are returned to higher temperatures, consistent with the complex having a role in establishing FLC repression.
Ben Trevaskis - One of the best experts on this subject based on the ideXlab platform.
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the central role of the vernalization1 gene in the vernalization response of cereals
Functional Plant Biology, 2010Co-Authors: Ben TrevaskisAbstract:Many varieties of wheat (Triticum spp.) and barley (Hordeum vulgare L.) require prolonged exposure to cold during winter in order to flower (vernalization). In these cereals, vernalization-induced flowering is controlled by the VERNALIZATION1 (VRN1) gene. VRN1 is a promoter of flowering that is activated by low temperatures. VRN1 transcript levels increase gradually during vernalization, with longer cold treatments inducing higher expression levels. Elevated VRN1 expression is maintained in the shoot apex and leaves after vernalization, and the level of VRN1 expression in these organs determines how rapidly vernalized plants flower. Some alleles of VRN1 are expressed without vernalization due to deletions or insertions within the promoter or first intron of the VRN1 gene. Varieties of wheat and barley with these alleles flower without vernalization and are grown where vernalization does not occur. The first intron of the VRN1 locus has histone modifications typically associated with the maintenance of an inactive chromatin state, suggesting this region is targeted by epigenetic mechanisms that contribute to repression of VRN1 before winter. Other mechanisms are likely to act elsewhere in the VRN1 gene to mediate low-temperature induction. This review examines how understanding the mechanisms that regulate VRN1 provides insights into the biology of vernalization-induced flowering in cereals and how this will contribute to future cereal breeding strategies.
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vernalization induced flowering in cereals is associated with changes in histone methylation at the vernalization1 gene
Proceedings of the National Academy of Sciences of the United States of America, 2009Co-Authors: Sandra N Oliver, E J Finnegan, W J Peacock, Elizabeth S Dennis, Ben TrevaskisAbstract:Prolonged exposure to low temperatures (vernalization) accelerates the transition to reproductive growth in many plant species, including the model plant Arabidopsis thaliana and the economically important cereal crops, wheat and barley. Vernalization-induced flowering is an epigenetic phenomenon. In Arabidopsis, stable down-regulation of FLOWERING LOCUS C (FLC) by vernalization is associated with changes in histone modifications at FLC chromatin. In cereals, the vernalization response is mediated by stable induction of the floral promoter VERNALIZATION1 (VRN1), which initiates reproductive development at the shoot apex. We show that in barley (Hordeum vulgare), repression of HvVRN1 before vernalization is associated with high levels of histone 3 lysine 27 trimethylation (H3K27me3) at HvVRN1 chromatin. Vernalization caused increased levels of histone 3 lysine 4 trimethylation (H3K4me3) and a loss of H3K27me3 at HvVRN1, suggesting that vernalization promotes an active chromatin state at VRN1. Levels of these histone modifications at 2 other flowering-time genes, VERNALIZATION2 and FLOWERING LOCUS T, were not altered by vernalization. Our study suggests that maintenance of an active chromatin state at VRN1 is likely to be the basis for epigenetic memory of vernalization in cereals. Thus, regulation of chromatin state is a feature of epigenetic memory of vernalization in Arabidopsis and the cereals; however, whereas vernalization-induced flowering in Arabidopsis is mediated by epigenetic regulation of the floral repressor FLC, this phenomenon in cereals is mediated by epigenetic regulation of the floral activator, VRN1.
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the molecular basis of vernalization induced flowering in cereals
Trends in Plant Science, 2007Co-Authors: Ben Trevaskis, Megan N Hemming, Elizabeth S Dennis, James W PeacockAbstract:Genetic analyses have identified three genes that control the vernalization requirement in wheat and barley; VRN1, VRN2 and FT (VRN3). These genes have now been isolated and shown to regulate not only the vernalization response but also the promotion of flowering by long days. VRN1 is induced by vernalization and accelerates the transition to reproductive development at the shoot apex. FT is induced by long days and further accelerates reproductive apex development. VRN2, a floral repressor, integrates vernalization and day-length responses by repressing FT until plants are vernalized. A comparison of flowering time pathways in cereals and Arabidopsis shows that the vernalization response is controlled by different MADS box genes, but integration of vernalization and long-day responses occurs through similar mechanisms.
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hvvrn2 responds to daylength whereas hvvrn1 is regulated by vernalization and developmental status
Plant Physiology, 2006Co-Authors: Ben Trevaskis, Megan N Hemming, James W Peacock, Elizabeth S DennisAbstract:Two genetic loci control the vernalization response in winter cereals; VRN1, which encodes an AP1-like MADS-box transcription factor, and VRN2, which has been mapped to a chromosome region containing ZCCT zinc finger transcription factor genes. We examined whether daylength regulates expression of HvVRN1 and HvVRN2. In a vernalization-responsive winter barley (Hordeum vulgare), expression of HvVRN1 is regulated by vernalization and by development, but not by daylength. Daylength affected HvVRN1 expression in only one of six vernalization-insensitive spring barleys examined and so cannot be a general feature of regulation of this gene. In contrast, daylength is the major determinant of expression levels of two ZCCT genes found at the barley VRN2 locus, HvZCCTa and HvZCCTb. In winter barley, high levels of HvZCCTa and HvZCCTb expression were detected only when plants were grown in long days. During vernalization in long-day conditions, HvVRN1 is induced and expression of HvZCCTb is repressed. During vernalization under short days, induction of HvVRN1 occurs without changes in HvZCCTa and HvZCCTb expression. Analysis of HvZCCTa and HvZCCTb expression levels in a doubled haploid population segregating for different vernalization and daylength requirements showed that HvVRN1 genotype determines HvZCCTa and HvZCCTb expression levels. We conclude that the vernalization response is mediated through HvVRN1, whereas HvZCCTa and HvZCCTb respond to daylength cues to repress flowering under long days in nonvernalized plants.
Caroline Dean - One of the best experts on this subject based on the ideXlab platform.
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the phd finger protein vrn5 functions in the epigenetic silencing of arabidopsis flc
Current Biology, 2007Co-Authors: Thomas Greb, Pedro Crevillen, Nuno Geraldo, Hailong An, Joshua S. Mylne, Anthony R. Gendall, Caroline DeanAbstract:Summary Vernalization, the acceleration of flowering by the prolonged cold of winter, ensures that plants flower in favorable spring conditions. During vernalization in Arabidopsis , cold temperatures repress FLOWERING LOCUS C ( FLC ) expression [1, 2] in a mechanism involving VERNALIZATION INSENSITIVE 3 (VIN3) [3], and this repression is epigenetically maintained by a Polycomb-like chromatin regulation involving VERNALIZATION 2 (VRN2), a Su(z)12 homolog, VERNALIZATION 1 (VRN1), and LIKE-HETEROCHROMATIN PROTEIN 1 [4–8]. In order to further elaborate how cold repression triggers epigenetic silencing, we have targeted mutations that result in FLC misexpression both at the end of the prolonged cold and after subsequent development. This identified VERNALIZATION 5 (VRN5), a PHD finger protein and homolog of VIN3. Our results suggest that during the prolonged cold, VRN5 and VIN3 form a heterodimer necessary for establishing the vernalization-induced chromatin modifications, histone deacetylation, and H3 lysine 27 trimethylation required for the epigenetic silencing of FLC . Double mutant and FLC misexpression analyses reveal additional VRN5 functions, both FLC -dependent and -independent, and indicate a spatial complexity to FLC epigenetic silencing with VRN5 acting as a common component in multiple pathways.
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variation in the epigenetic silencing of flc contributes to natural variation in arabidopsis vernalization response
Genes & Development, 2006Co-Authors: Chikako Shindo, Pedro Crevillen, Clare Lister, Magnus Nordborg, Caroline DeanAbstract:Vernalization, the cold-induced acceleration of flowering, involves the epigenetic silencing of the floral repressor gene FLOWERING LOCUS C (FLC). We investigated the molecular basis for variation in vernalization in Arabidopsis natural accessions adapted to different climates. A major variable was the degree to which different periods of cold caused stable FLC silencing. In accessions requiring long vernalization, FLC expression was reactivated following nonsaturating vernalization, but this reactivation was progressively attenuated with increasing cold exposure. This response was correlated with the rate of accumulation of FLC histone H3 Lys 27 trimethylation (H3K27me3). Thus, variation in epigenetic silencing of FLC appears to have contributed to Arabidopsis adaptation.
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multiple roles of arabidopsis vrn1 in vernalization and flowering time control
Science, 2002Co-Authors: Yaron Y Levy, Joshua S. Mylne, Stephane Mesnage, Anthony R. Gendall, Caroline DeanAbstract:Arabidopsis VRN genes mediate vernalization, the process by which a long period of cold induces a mitotically stable state that leads to accelerated flowering during later development. VRN1 encodes a protein that binds DNA in vitro in a non–sequence-specific manner and functions in stable repression of the major target of the vernalization pathway, the floral repressor FLC . Overexpression of VRN1 reveals a vernalization-independent function for VRN1 , mediated predominantly through the floral pathway integrator FT , and demonstrates that VRN1 requires vernalization-specific factors to target FLC .
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the vernalization 2 gene mediates the epigenetic regulation of vernalization in arabidopsis
Cell, 2001Co-Authors: Anthony R. Gendall, Yaron Y Levy, Allison Wilson, Caroline DeanAbstract:Abstract The acceleration of flowering by a long period of low temperature, vernalization, is an adaptation that ensures plants overwinter before flowering. Vernalization induces a developmental state that is mitotically stable, suggesting that it may have an epigenetic basis. The VERNALIZATION2 ( VRN2 ) gene mediates vernalization and encodes a nuclear-localized zinc finger protein with similarity to Polycomb group (PcG) proteins of plants and animals. In wild-type Arabidopsis , vernalization results in the stable reduction of the levels of the floral repressor FLC . In vrn2 mutants, FLC expression is downregulated normally in response to vernalization, but instead of remaining low, FLC mRNA levels increase when plants are returned to normal temperatures. VRN2 function therefore stably maintains FLC repression after a cold treatment, serving as a mechanism for the cellular memory of vernalization.
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arabidopsis mutants showing an altered response to vernalization
Plant Journal, 1996Co-Authors: John Chandler, Allison Wilson, Caroline DeanAbstract:Flowering in many plant species is accelerated by a long period of cold temperature, known as a vernalization period. This research investigates how this cold temperature signal is perceived by plant cells and the mechanism by which it influences the transition to flowering. Mutagenesis of the late-flowering, vernalization-responsive, Arabidopsis mutant, fca, has yielded five independent mutations (termed vrn mutations) conferring an altered vernalization response. Allelism tests showed that these mutations fall into at least three complementation groups defining three loci named VRN 1, 2 and 3. The vrn1 and vrn2 mutations did not affect the acclimation response as judged by expression of cold-induced transcripts and freezing tolerance assays. vrn1-1 affected the short-day vernalization response of Landsberg erecta and reduced the vernalization response of other late-flowering Arabidopsis mutants. The acceleration of flowering by GA3 was not affected by vrn1-1. The VRN 1 locus was mapped to chromosome 3.
Ildiko Karsai - One of the best experts on this subject based on the ideXlab platform.
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validation of the vrn h2 vrn h1 epistatic model in barley reveals that intron length variation in vrn h1 may account for a continuum of vernalization sensitivity
Molecular Genetics and Genomics, 2007Co-Authors: Peter Szucs, Tony H H Chen, Alfonso Cuestamarcos, Kale G Haggard, Ann Corey, Ildiko Karsai, Jeffrey S Skinner, Patrick M HayesAbstract:The epistatic interaction of alleles at the VRN-H1 and VRN-H2 loci determines vernalization sensitivity in barley. To validate the current molecular model for the two-locus epistasis, we crossed homozygous vernalization-insensitive plants harboring a predicted “winter type” allele at either VRN-H1 (Dicktoo) or VRN-H2 (Oregon Wolfe Barley Dominant), or at both VRN-H (Calicuchima-sib) loci and measured the flowering time of unvernalized F2 progeny under long-day photoperiod. We assessed whether the spring growth habit of Calicuchima-sib is an exception to the two-locus epistatic model or contains novel “spring” alleles at VRN-H1 (HvBM5A) and/or VRN-H2 (ZCCT-H) by determining allele sequence variants at these loci and their effects relative to growth habit. We found that (a) progeny with predicted “winter type” alleles at both VRN-H1 and VRN-H2 alleles exhibited an extremely delayed flowering (i.e. vernalization-sensitive) phenotype in two out of the three F2 populations, (b) sequence flanking the vernalization critical region of HvBM5A intron 1 likely influences degree of vernalization sensitivity, (c) a winter habit is retained when ZCCT-Ha has been deleted, and (d) the ZCCT-H genes have higher levels of allelic polymorphism than other winterhardiness regulatory genes. Our results validate the model explaining the epistatic interaction of VRN-H2 and VRN-H1 under long-day conditions, demonstrate recovery of vernalization-sensitive progeny from crosses of vernalization-insensitive genotypes, show that intron length variation in VRN-H1 may account for a continuum of vernalization sensitivity, and provide molecular markers that are accurate predictors of “winter vs spring type” alleles at the VRN-H loci.
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validation of the two gene epistatic model for vernalization response in a winter spring barley cross
Euphytica, 2006Co-Authors: K Koti, P Szűcs, G Kiss, Katalin Mészáros, Cs Horvath, Zoltan Bedő, Ildiko Karsai, P M HayesAbstract:A two gene epistatic model in which a dominant “winter growth habit” allele at Vrn-H2 encodes a repressor with a corresponding binding site in a recessive vrn-H1 allele explains the vernalization response phenotypes in an array of barley germplasm. In order to validate the model genetically, we developed an F 2 population (and F 2-derived F 3 families) from the cross of Hardy (winter) × Jubilant (spring). Using gene-specific primers, we determined the Vrn-H1 and Vrn-H2 allele architecture of each F 2 plant and we measured the growth habit phenotype of each F 2 plant via phenotyping of its F 3 progeny under controlled environment conditions. We used a set of treatments involving plus/minus vernalization under long photoperiod and vernalization under short photoperiod. Alleles at the two loci showed expected patterns of segregation and independent assortment. Under long day conditions, the two Vrn genes were the primary determinants of heading date, regardless of the vernalization treatment. Under short photoperiod, the effects of these loci were not significant. There was incomplete dominance at Vrn-H1: heterozygotes were significantly later to head than Vrn-H1Vrn-H1 genotypes. Vrn-H2 genotypes were also significantly later to head, even when plants were vernalized. These results validate the two-gene epistatic model for vernalization response under long-day conditions. The results under short photoperiod, and the variance in flowering with vernalization, confirm that while the two Vrn genes are the primary determinants of vernalization response, they are part of a larger interactome that determines the timing of the vegetative to reproductive transition.
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positional relationships between photoperiod response qtl and photoreceptor and vernalization genes in barley
Theoretical and Applied Genetics, 2006Co-Authors: Peter Szucs, L L D Cooper, Yong Q Gu, Tony H H Chen, Patrick M Hayes, Joachim Von Zitzewitz, Katalin Mészáros, Ildiko Karsai, Jeffrey S SkinnerAbstract:Winterhardiness has three primary components: photoperiod (day length) sensitivity, vernalization response, and low temperature tolerance. Photoperiod and vernalization regulate the vegetative to reproductive phase transition, and photoperiod regulates expression of key vernalization genes. Using two barley mapping populations, we mapped six individual photoperiod response QTL and determined their positional relationship to the phytochrome and cryptochrome photoreceptor gene families and the vernalization regulatory genes HvBM5A, ZCCT-H, and HvVRT-2. Of the six photoreceptors mapped in the current study (HvPhyA and HvPhyB to 4HS, HvPhyC to 5HL, HvCry1a and HvCry2 to 6HS, and HvCry1b to 2HL), only HvPhyC coincided with a photoperiod response QTL. We recently mapped the candidate genes for the 5HL VRN-H1 (HvBM5A) and 4HL VRN-H2 (ZCCT-H) loci, and in this study, we mapped HvVRT-2, the barley TaVRT-2 ortholog (a wheat flowering repressor regulated by vernalization and photoperiod) to 7HS. Each of these three vernalization genes is located in chromosome regions determining small photoperiod response QTL effects. HvBM5A and HvPhyC are closely linked on 5HL and therefore are currently both positional candidates for the same photoperiod effect. The coincidence of photoperiod-responsive vernalization genes with photoperiod QTL suggests vernalization genes should also be considered candidates for photoperiod effects.
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the vrn h2 locus is a major determinant of flowering time in a facultative winter growth habit barley hordeum vulgare l mapping population
Theoretical and Applied Genetics, 2005Co-Authors: Ildiko Karsai, T Filichkina, P M Hayes, L Lang, Peter Szucs, Katalin Mészáros, Jeffrey S Skinner, Zoltán BedoAbstract: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.
