拟南芥隐花色素相关基因功能分析
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摘要
隐花色素(CRYPTOCHROME,简称CRY)是与DNA光解酶氨基酸序列高度同源的黄素蛋白,但不具有光解酶活性。拟南芥的隐花色素家族有三个成员,其中隐花色素1(CRY1)主要抑制下胚轴伸长,而隐花色素2(CRY2)主要调节光周期开花,并且这两个光受体具有部分冗余的功能。为进一步阐明拟南芥隐花色素的生物学功能,本实验室采用酵母双杂交的方法,以GUS-CCT2为诱饵蛋白筛选拟南芥cDNA文库,筛选到与之相互作用的蛋白RAT1(Related to acyltransferase)。rat1单突变体在长日条件下(16 h光照/8 h黑暗)开花时间与其野生型一致。在sgs3-11突变体中过量表达RAT1的拟南芥转基因植株,在长日条件下开花时间与其野生型一致。共聚焦激光扫描显微镜结果表明GFP-RAT1在细胞质和核中均有表达。通过序列比对发现:RAT2与RAT1的氨基酸序列同源性达89.7%,与γ-型碳酸酐酶基因在同一个进化枝上。rat2单突变体在长日条件下开花时间与其野生型一致。我们试图构建rat1rat2双突变体,结果发现不能得到rat1/-rat2/-双纯合的突变体。亚历山大染色结果发现半杂合的rat1/+ rat2/-突变体的花粉正常。但其果荚中约有23.7%的胚是败育的。胚经透明之后,在微分干涉相差显微镜下观察发现rat1/+ rat2/-突变体的中胚发育推迟,最终不能发育成正常的胚,说明RAT1和RAT2基因可能参与胚发育的调节。我们又构建了RAT1-RNAi载体,转化rat2单突变体。转基因植株r2r1i(rat2/RAT1-RNAi)在长日条件下表现出晚开花表型。实时定量PCR研究结果表明,转基因植株r2r1i晚开花表型可能与开花基因FT的下调和FLC的上调有关。转基因植株r2r1i对光敏感,其下胚轴在不同的光下均比野生型变短,说明RAT1和RAT2基因可能以功能冗余的方式参与光形态建成。将RAT2与YFP融合转化拟南芥原生质体。共聚焦激光扫描显微镜结果表明RAT2-YFP定位于线粒体,说明RAT2也有可能在线粒体中起作用。
cry1cry2双突变体在长日条件下比野生型晚开花。采用激活标签的方法,在cry1cry2双突变体背景下筛选到一个晚开花突变体ecc2 (enhencer of cry1cry2)。TAIL-PCR结果表明,ecc2突变体的T-DNA插入在第二条染色体上At2g45430开放阅读框附近。ecc2突变体晚开花的表型很可能是由于AHL22 (At2g45430)基因的过量表达引起的。凝胶电泳迁移率实验(EMSA)结果表明AHL22可以结合到富含AT碱基的DNA序列上,说明AHL22是一个AT-hook蛋白。采用荧光显微镜观察GFP-AHL22的亚细胞定位发现AHL22定位于细胞核。将野生型植株与GFP-AHL22转基因植株的细胞核纯化出来之后,采用免疫印迹(Western blot)的方法检测发现AHL22在细胞核中表达。进一步研究发现GFP-AHL22蛋白不受不同光的调节。我们的研究还发现AHL22蛋白丰度在根中较高,说明AHL22有可能在根中起作用。采用根特异性的启动子TobRB7驱动GFP-AHL22在根中表达,结果表明TobRB7:GFP-AHL22转基因植株开花时间比野生型略晚。共聚焦激光扫描显微镜结果表明只在TobRB7:GFP-AHL22转基因植株的根中观察到GFP荧光。
此外,还筛选到另一个早花激活标签突变体,scc17-D (suppressor of cry1cry2#17-Dominant)。TAIL-PCR结果表明,scc17-D突变体的T-DNA插入在第一条染色体上的At1g25440和At1g25450两个基因之间。scc17-D突变体表现出植株矮化、叶片变小、果荚变小和子叶表皮细胞形状发生改变等性状。结果表明,scc17-D突变体对植物的发育具有广泛的调节功能。本研究为拟南芥隐花色素的功能研究提供了一个新的突变体。
Cryptochromes are flavoproteins that share similarity in the sequence of amino acids to DNA photolyase but lack the photolyase activity. There are three members of the cryptochromes gene family in Arabidopsis thaliana. Cryptochrome1 (CRY1) and CRY2 mediate primarily blue light inhibition of hypocotyl elongation and photoperiodic promotion of floral initiation, respectively, and they functions redundantly. In order to further study the biological functions of Arabidopsis thaliana cryptochromes, using GUS-CCT2 as a bait, we screened the cDNA library and found that RAT1 (Related to acyltransferase) can interact with GUS-CCT2. The rat1 single mutant flowered at nearly the same time as wild-type under long day conditions (16 h light/8 h dark). We overexpressed RAT1 in the sgs3-11 mutant and found that the RAT1-ox transgenic lines flowered at the similar time as the wild-type. A fusion protein consisting of the green fluorescent protein (GFP) and RAT1 was expressed in Arabidopsis and the localization of the fusion proteins was determined by confocal laser scanning microscopy. The results showed that GFP-RAT1 was localized to not only the cytoplasm and but also the nucleus. Sequences alignment showed that RAT2 shared 89.7% amino acids sequences similarity with RAT1, which were both in the same clade as carbonic anhydrases genes. The rat2 single mutant flowered also at the similar time as wild-type plants under long day conditions. We tried to construct rat1/-rat2/- double mutants and found that the double homozygous rat1/-rat2/- mutants were lethal. Alexander staining showed that the pollen’s activity was normal in the heterozygous rat1/+ rat2/- double mutants. About 23.7% of embryos of rat1/+ rat2/- double mutants showed defects. The aborted embryos were cleared and observed under differential interference contrast (DIC) microscopy, showing that they developed slower than those of normal embryos and were incapable of becoming fully developed. This indicated that RAT1 and RAT2 might involve in embryogenesis. We transformed rat2 with the RAT1-RNAi construct. The transgenic r2r1i (rat2/RAT1-RNAi) lines flowered later than that of wild type under long day conditions. Real time Q-PCR results showed that the late flowering phenotype of r2r1i lines might result from the downregulation of Flowering Locus T (FT) and upregulation of Flowering Locus C (FLC). The r2r1i lines were hypersensitive to light and their hypocotyls were shorter than those of wild type under different light conditions, which indicated that RAT1 and RAT2 might act redundantly in photomorphogenesis. We fused RAT2 to yellow fluorescent protein (YFP) and transformed Arabidopsis protoplasts with RAT2-YFP. The localization of RAT2-YFP was determined by confocal laser scanning microscopy, and it showed that RAT2-YFP was localized to the mitochondria. This indicated that RAT2 might play a role in mitochondria.
The cry1cry2 double mutant flowered later than wild-type under long day conditions. Through activation tagging, we identified a late flowering mutant ecc2 (enhencer of cry1cry2). TAIL-PCR results showed that a T-DNA insertion located near the open reading frame of At2g45430 on the second chromosome, suggesting that the late flowering phenotype of ecc2 mutant might be caused by overexpression of AHL22 (At2g45430) gene. The electrophoretic mobility shift assay (EMSA) showed that AHL22 can bind specifically to AT-rich DNA sequence, indicating that AHL22 is a bona fide AT-hook protein. The subcellular localization of GFP-AHL22 was observed in the nucleus. The nuclear proteins were purified from seedlings of wild type plants and GFP-AHL22 transgenic lines, and the western blot results showed that AHL22 was expressed in the nucleus. Additionally, GFP-AHL22 was not regulated by light. We also found that AHL protein abundance was higher in roots, indicating that AHL22 might play a role in roots. Then we constructed a GFP-AHL22 fusion gene drived by a root specific promoter TobRB7. The TobRB7:GFP-AHL22 transgenic lines flowered slightly later than that of wild type plants. The GFP-AHL22 fluorescence signal was only observed in the roots of TobRB7:GFP-AHL22 transgenic lines through confocal laser scanning microscopy.
Through activation tagging, we identified an early flowering mutant called scc17-D (suppressor of cry1cry2#17-Dominant). TAIL-PCR results showed that a T-DNA insertion located between At1g25440 and At1g25450 on the first chromosome. The scc17-D mutant displayed phenotypes of dwarfism, reduction in leaf size, reduced length of siliques, and the altered shape of cotyledon pavement cells, suggesting that scc17-D had extensive functions on plant development. The results provide a new mutant for genetic studies of cryptochrome functions in Arabidopsis thaliana.
cry1cry2双突变体在长日条件下比野生型晚开花。采用激活标签的方法,在cry1cry2双突变体背景下筛选到一个晚开花突变体ecc2 (enhencer of cry1cry2)。TAIL-PCR结果表明,ecc2突变体的T-DNA插入在第二条染色体上At2g45430开放阅读框附近。ecc2突变体晚开花的表型很可能是由于AHL22 (At2g45430)基因的过量表达引起的。凝胶电泳迁移率实验(EMSA)结果表明AHL22可以结合到富含AT碱基的DNA序列上,说明AHL22是一个AT-hook蛋白。采用荧光显微镜观察GFP-AHL22的亚细胞定位发现AHL22定位于细胞核。将野生型植株与GFP-AHL22转基因植株的细胞核纯化出来之后,采用免疫印迹(Western blot)的方法检测发现AHL22在细胞核中表达。进一步研究发现GFP-AHL22蛋白不受不同光的调节。我们的研究还发现AHL22蛋白丰度在根中较高,说明AHL22有可能在根中起作用。采用根特异性的启动子TobRB7驱动GFP-AHL22在根中表达,结果表明TobRB7:GFP-AHL22转基因植株开花时间比野生型略晚。共聚焦激光扫描显微镜结果表明只在TobRB7:GFP-AHL22转基因植株的根中观察到GFP荧光。
此外,还筛选到另一个早花激活标签突变体,scc17-D (suppressor of cry1cry2#17-Dominant)。TAIL-PCR结果表明,scc17-D突变体的T-DNA插入在第一条染色体上的At1g25440和At1g25450两个基因之间。scc17-D突变体表现出植株矮化、叶片变小、果荚变小和子叶表皮细胞形状发生改变等性状。结果表明,scc17-D突变体对植物的发育具有广泛的调节功能。本研究为拟南芥隐花色素的功能研究提供了一个新的突变体。
Cryptochromes are flavoproteins that share similarity in the sequence of amino acids to DNA photolyase but lack the photolyase activity. There are three members of the cryptochromes gene family in Arabidopsis thaliana. Cryptochrome1 (CRY1) and CRY2 mediate primarily blue light inhibition of hypocotyl elongation and photoperiodic promotion of floral initiation, respectively, and they functions redundantly. In order to further study the biological functions of Arabidopsis thaliana cryptochromes, using GUS-CCT2 as a bait, we screened the cDNA library and found that RAT1 (Related to acyltransferase) can interact with GUS-CCT2. The rat1 single mutant flowered at nearly the same time as wild-type under long day conditions (16 h light/8 h dark). We overexpressed RAT1 in the sgs3-11 mutant and found that the RAT1-ox transgenic lines flowered at the similar time as the wild-type. A fusion protein consisting of the green fluorescent protein (GFP) and RAT1 was expressed in Arabidopsis and the localization of the fusion proteins was determined by confocal laser scanning microscopy. The results showed that GFP-RAT1 was localized to not only the cytoplasm and but also the nucleus. Sequences alignment showed that RAT2 shared 89.7% amino acids sequences similarity with RAT1, which were both in the same clade as carbonic anhydrases genes. The rat2 single mutant flowered also at the similar time as wild-type plants under long day conditions. We tried to construct rat1/-rat2/- double mutants and found that the double homozygous rat1/-rat2/- mutants were lethal. Alexander staining showed that the pollen’s activity was normal in the heterozygous rat1/+ rat2/- double mutants. About 23.7% of embryos of rat1/+ rat2/- double mutants showed defects. The aborted embryos were cleared and observed under differential interference contrast (DIC) microscopy, showing that they developed slower than those of normal embryos and were incapable of becoming fully developed. This indicated that RAT1 and RAT2 might involve in embryogenesis. We transformed rat2 with the RAT1-RNAi construct. The transgenic r2r1i (rat2/RAT1-RNAi) lines flowered later than that of wild type under long day conditions. Real time Q-PCR results showed that the late flowering phenotype of r2r1i lines might result from the downregulation of Flowering Locus T (FT) and upregulation of Flowering Locus C (FLC). The r2r1i lines were hypersensitive to light and their hypocotyls were shorter than those of wild type under different light conditions, which indicated that RAT1 and RAT2 might act redundantly in photomorphogenesis. We fused RAT2 to yellow fluorescent protein (YFP) and transformed Arabidopsis protoplasts with RAT2-YFP. The localization of RAT2-YFP was determined by confocal laser scanning microscopy, and it showed that RAT2-YFP was localized to the mitochondria. This indicated that RAT2 might play a role in mitochondria.
The cry1cry2 double mutant flowered later than wild-type under long day conditions. Through activation tagging, we identified a late flowering mutant ecc2 (enhencer of cry1cry2). TAIL-PCR results showed that a T-DNA insertion located near the open reading frame of At2g45430 on the second chromosome, suggesting that the late flowering phenotype of ecc2 mutant might be caused by overexpression of AHL22 (At2g45430) gene. The electrophoretic mobility shift assay (EMSA) showed that AHL22 can bind specifically to AT-rich DNA sequence, indicating that AHL22 is a bona fide AT-hook protein. The subcellular localization of GFP-AHL22 was observed in the nucleus. The nuclear proteins were purified from seedlings of wild type plants and GFP-AHL22 transgenic lines, and the western blot results showed that AHL22 was expressed in the nucleus. Additionally, GFP-AHL22 was not regulated by light. We also found that AHL protein abundance was higher in roots, indicating that AHL22 might play a role in roots. Then we constructed a GFP-AHL22 fusion gene drived by a root specific promoter TobRB7. The TobRB7:GFP-AHL22 transgenic lines flowered slightly later than that of wild type plants. The GFP-AHL22 fluorescence signal was only observed in the roots of TobRB7:GFP-AHL22 transgenic lines through confocal laser scanning microscopy.
Through activation tagging, we identified an early flowering mutant called scc17-D (suppressor of cry1cry2#17-Dominant). TAIL-PCR results showed that a T-DNA insertion located between At1g25440 and At1g25450 on the first chromosome. The scc17-D mutant displayed phenotypes of dwarfism, reduction in leaf size, reduced length of siliques, and the altered shape of cotyledon pavement cells, suggesting that scc17-D had extensive functions on plant development. The results provide a new mutant for genetic studies of cryptochrome functions in Arabidopsis thaliana.
引文
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