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Brassica rapa種の開花時期の遺伝的制御

AYASHA AKTER 神戸大学

2020.09.25

概要

Brassica is an important genus in the family of Brassicaceae which includes oilseeds, condiments and vegetables. Brassica species undergoes a whole genome triplication event compared with the model plant Arabidopsis thaliana that plays an important role in the speciation and wide morphotype diversification of Brassica. Diploid species such as Brassica rapa and Brassica oleracea have three copies of genes orthologous to each A. thaliana gene, though deletion in one or two of the three homologs has occurred in some genes after the whole genome triplication. The floral transition is one of the crucial events in plant's life history; flowering time is an important agricultural trait. Flowering time is also important for the yield of crops or vegetables, and the regulation of flowering time is an important goal of plant breeding. There is a wide variation in flowering time within species of the genus Brassica, and this variation is largely dependent on a difference of vernalization requirements. Vernalization is exposure to prolonged cold that alters gene expression and accelerates a transition from the vegetative to reproductive phase. Premature flowering induced by continuous low temperature critically decreases the yield and quality of the harvested product. Therefore, high bolting resistance is an important breeding trait, and understanding the molecular mechanism of vernalization is necessary to achieve this goal. In A. thaliana a relative of B. rapa, two genes, FRIG1DA (FRT) and FLOWERING LOCUS C (FLC) are important in regulating flowering time. The vernalization response including the repression of FLC expression by cold treatment is similar to A. thaliana. B. rapa and B. oleracea each have four homologs of FLC, and the allotetraploid species, Brassica napus, has nine homologs. The increased number of homologs makes their role in vernalization more complicated; among homologs in an individual plant, there are variations in the transcriptional levels of FLC before and after vernalization, in addition to variation of FLC levels between lines. During my study, I demonstrated that BrFR1b functions as an activator of BrFLC which repress flowering in B. rapa. I also confirmed that BrFLC1, BrFLC2, and BrFLC3 act as floral repressors by producing transgenic plants overexpressing each FLC gene and finding a delay in flowering. I showed a positive correlation between the steady state expression levels of the sum of the BrFLC paralogs and the days to flowering after four weeks of cold treatment, suggesting that this is an indicator of the vernalization requirement. As histone modifications can be important for controlling expression levels of genes, I compared the level of the repressive histone mark H3K27me3 between lines, between tissues, or between before and after vernalization by chromatin immunoprecipitation sequencing (ChIP-seq) at whole genome levels in B. rapa. About 90% of the same genes had H3K27me 3 in the two B. rapa lines examined, while there is greater variation between different tissues, indicating that H3K27me3 distribution is conserved between lines but differs between tissues within a line. I also confirmed that genes having H3K27me3 marks showed lower expression levels and tissue -specific expression .1 examined H3K27me 3 states in non-vernalized, vernalized, and after return to warm condition following vernalization, and most of the gene had H3K27me3 levels unchanged by cold treatment. Furthermore, I identified that increased H3K27me3 following vernalization included four BrFLC genes where H3K27me3 was enriched around the transcription start sites. Following return to normal temperature growth conditions, H3K27me3 spread along all four BrFLC paralogs providing stable repression of the gene. This is the important information about H3K27me 3 states at whole genome level and H3K27me3 states after vernalization in B. rapa, suggests that a PHD-PRC2 complex that contains the enzyme responsible for the histone H3K27me3 is recruited to the gene regions of the BrFLCs, which induces stable epigenetic repression .1 also performed ChIP-seq experiments showed that a few genes showed spreading of H3K27me 3 like in the case in BrFLCs, suggesting that this epigenetic change in vernalization is specific to only a few genes including four BrFLC genes. I also observed the differential gene expression pattern during vernalization in B. rapa. I did the transcriptome analysis with different durations of vernalization treatments in B. rapa and found most genes were downregulated following cold treatment. BrFLC expression decreased and expression of genes known to transmit the vernalization state BrVIN3 and BrSOC1 were induced during cold treatment. During cold treatment, some flowering pathway genes showed different expression patterns between paralogs and some genes whose orthologous genes in A. thaliana were unchanged showed change of expression in B. rapa. I suggested that Chinese cabbage’s flowering pathway is different and more complex than in A. thaliana. These outcomes provide significant insights into the genetic control of bolting and flowering that occurred during the vernalization in B. rapa.

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