黄梁木α扩展蛋白的基因克隆及功能研究
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摘要
黄梁木(Anthocephalus chinensis)别名团花树,属茜草科团花属常绿阔叶乔木,是亚洲、南亚热带的速生树种。由于其速生性,在1972年的世界林业大会上被公认为“奇迹树”,在我国华南地区有着巨大的开发和应用前景。然而,由于黄梁木遗传信息十分匮乏,其木材速生性的分子生物学基础研究至今尚未开展。扩展蛋白作为一类细胞壁生长重要调控因子,与植物的生长发育息息相关。因此,本文以黄梁木的α扩展蛋白编码基因EXPA为切入点,筛选出其16个家族成员中在形成层高丰度表达的基因并进行了功能鉴定。首先,利用简并引物扩增同源基因保守序列,结合基因组步移及3'RACE技术克隆到扩展蛋白基因序列并进行生物信息学分析。在此基础上,筛选合适内参基因,对α扩展蛋白基因在黄梁木中的时空表达进行分析。选择形成层中高表达的AcEXPA8基因进行基因组中的拷贝数分析,并进行蛋白质三维结构建模预测其功能。同时,通过构建植物表达载体并转化拟南芥研究,进一步鉴定AcEXPA8基因的功能。通过上述研究,得到主要结果如下：
1.通过设计保守区的简并引物,PCR扩增获得同源基因保守序列,再结合基因组步移和3'RACE技术获得黄梁木EXPA基因家族的16个成员,分别命名为AcEXPAl-16, GenBank注册号为FJ417847、JF922686-JF922700,其相对应的基因组DNA序列的注册号为GQ228823、JF922701-JF922715。
2.黄梁木AcEXPAl-16基因的内含子分布方式一致,都含有EXPA基因保守的内含子1和内含子2。除了AcEXPA4中的内含子2的剪接方式是TA-GA,其余都符合真核生物内含子GT-AG的剪接模式。
3.AcEXPAl-16基因编码的氨基酸序列都含有EXPA蛋白保守序列,包括N端的信号肽、8个C残基、1个组氨酸域(His-Phe-Asp, HFD),和羧基末端的4个保守色氨酸(tryptophan, W)残基,但在AcEXPA12的第一个W被亮氨酸(leucine, L)所取代。在AcEXPA1、AcEXPA5、AcEXPA8和AcEXPA9成熟蛋白的N端和C端都各自含有RIPGV和KNFRV序列。并通过系统进化分析,黄梁木AcEXPA家族成员分别属于4个亚家族。
4.筛选到环孢素A受体基因AcCyP作为内参基因用于荧光定量PCR分析。AcEXPA1-16基因在黄梁木中具有明显的组织表达特异性：在四个幼嫩的生长组织中,AcEXPA8在形成层的表达量最高,AcEXPA4、AcEXPA6、AcEXPA15、AcEXPA14和AcEXPA10在嫩叶中大量表达,AcEXPA13只在嫩叶中有所表达,而AcEXPA16仅在嫩根中表达。AcEXPA8在不同生长季节的形成层中的表达也明显不同,7月的最高,1月的最低。在维管发育径向的不同细胞层,AcEXPA8在次生壁形成区的表达量明显低于形成层。
5. AcEXPA8基因以多拷贝的形式在黄梁木基因组中存在。蛋白同源建模显示成熟AcEXPA8蛋白的三维结构与到目前为止仅知的EXP蛋白玉米ZmEXPB1晶体结构极为相似,含有domain1和domain2以及两者之间的连接区域。
6.通过转AcEXPA8基因拟南芥研究,转基因植株与野生型相比,叶片形成加速,叶片面积增大,并且叶片长生长大于宽生长,使得叶片形状变得扁长,在基叶完全成熟期,基座的半径明显增大。而且,转基因植株的高度和茎的粗度也明显大于野生型。
7.过表达AcEXPA8基因促进了拟南芥的次生生长,主要是导致了纤维细胞增长和细胞壁的加厚,但没有改变细胞壁中纤维素的含量。此外,转基因植株茎的髓和皮层薄壁细胞明显增大,而皮层的薄壁细胞在沿周长方向的长度增长尤为明显。
本研究克隆到并分析了黄梁木AcEXPA基因家族成员在不同幼嫩组织的表达差异,对形成层中高丰度表达的AcEXPA8基因的时空表达进行了进一步分析并进行了转基因功能鉴定,结果为今后研究α扩展蛋白基因与团花树木质部生长速度和材质的关系提供基础,并最终为林木分子育种提供潜在的候选基因。
Anthocephalus chinensis with the synonym of Neolamarckia cadamba belonging to the family Rubiaceae, is an evergreen broadleaf arbor and a fast-growing tree species which grows in Tropical Asia and Southern Asia. It was universally accepted as "a miraculous tree" in the World Forestry Congress in1972because of its rapid growth. Meanwhile, it possesses significant development and utilization value in South China. However, genetic information on A. chinensis is still unclear, especially in molecular mechanism study on fast-growing of wood. Expansin is a class of important cell growth regulator, and closely related to the growth and development of plants. So, in the study, we fitstly cloned a-expansin gene sequences in A. chinensis, and inditified the function of the a-expansin gene abundantly expressed in the cambium region, a-expansin genes were isolated by amplifying conservative domain binding with genomic walking and3'RACE techniques from four differential growth tissues in A. chinensis, followed by bioinformatic analysis. Based on these, we selected a suitable reference gene and analysed the temporal and spatial expression pattern of a-expansin genes. AcEXPA8gene expressed highly in cambium region was selected to analyse its copy number in genome, and we also predicted the protein structure and function by homology modeling. Meanwhile, we constructed the plant expression vector and transformed it into Arabidopsis thaliana to further indentify AcEXPA8gene function. The main results and conclusions obtained in this study are shown as follows:
1. AcEXPA1-16genes were cloned by a cloning strategy. Firstly, the conservative regions of A. chinensis a-expansin genes were amplified with the degenerated primers, then, the flank sequences were obtained by genomic walking and3'RACE. Their GenBank accession Nos. were FJ417847, JF922686-JF922700, with corresponding genomic DNA sequences of GenBank accession Nos. GQ228823, JF922701-JF922715.
2. The intron-exon structures of AcEXPAl-16are similar between them, and they all contain intronl and intron2which are conservative in a-expansin genes. All AcEXPAl-16introns conforn with eukaryotic intron splicing pattern of GT-AG, except the intron2of AcEXPA4with TA-GA.
3. All amino acid sequences encoded by AcEXPAI-16gene contain conservative sequences of EXPA protein, including the N-terminal signal peptide, eight C residues, HFD motif and four W residues near the C terminus. But, there is a leucine (L) instead of the first W in AcEXPA12. All mature proteins of AcEXPA1, AcEXPA5, AcEXPA8and AcEXPA9cotain RIPGV sequence at N-terminal and KNFRV sequence at C-terminal. And AcEXPAl-16genes were divided into four subfamilies in the phylogenetic analysis.
4. The screened cyclophilin gene AcCyP was suitable in RT-qPCR analysis as reference gene. AcEXPA1-16genes showed obviously tissue-specific expression in A. chinensis. In the four young tissues in growth, AcEXPA8was expressed highestly in cambium region. AcEXPA4, AcEXPA6, AcEXPA15, AcEXPA14and AcEXPA10were abundantly expressed in the tender leaf. AcEXPA13was only expressed in leaf, whereas AcEXPA16was only expressed in root. The expression of AcEXPA8was significantly different between different growing seasons in cambium region, highest in July, but lowest in January. And the abundance of AcEXPA8expressed in second wall formation region was significantly lower than that in camium region.
5. AcEXPA8gene is present in A. chinensis genomic DNA as multicopy. Protein homology modeling showed that the3D structure of mature protein AcEXPA8is similar to maize ZmEXPB1which is the only known crystal structure of EXP protein so far, and also contains domain1, domain2and connection area between them.
6. Overexpression of AcEXPA8in Arabidopsis could not only accelerate leaf formation, induce leaf area, cause leaf length growth greater than width growth, leading to the leaf shape becoming prolate, and increase the radius of fully mature rosette, but also increase plant height and stem diameter.
7. In transgenic plant stem, overexpression of AcEXPA8promoted secondary growth in Arabidopsis, mainly leading to increase fiber cell length and cell wall thickness, but without changing the content of cellulose in the cell wall. In addition, the area of pith and cortex parenchyma cells increased obviously, and the length of cortex parenchyma cells along the circumference direction increased particularly.
In this study, the AcEXPA gene family member expressions were analyzed in different young tissures, and the temporal and spatial expression pattern of AcEXPA8gene expressed abundantly in cambium region was further analized, followed by gene function analysis. These findings will lay the foundation for studying the relationship between a-expansin genes and growth rate and wood quality of the xylem in A. chinensis, and finally provide potential candidate genes for tree molecular breeding.
1.通过设计保守区的简并引物,PCR扩增获得同源基因保守序列,再结合基因组步移和3'RACE技术获得黄梁木EXPA基因家族的16个成员,分别命名为AcEXPAl-16, GenBank注册号为FJ417847、JF922686-JF922700,其相对应的基因组DNA序列的注册号为GQ228823、JF922701-JF922715。
2.黄梁木AcEXPAl-16基因的内含子分布方式一致,都含有EXPA基因保守的内含子1和内含子2。除了AcEXPA4中的内含子2的剪接方式是TA-GA,其余都符合真核生物内含子GT-AG的剪接模式。
3.AcEXPAl-16基因编码的氨基酸序列都含有EXPA蛋白保守序列,包括N端的信号肽、8个C残基、1个组氨酸域(His-Phe-Asp, HFD),和羧基末端的4个保守色氨酸(tryptophan, W)残基,但在AcEXPA12的第一个W被亮氨酸(leucine, L)所取代。在AcEXPA1、AcEXPA5、AcEXPA8和AcEXPA9成熟蛋白的N端和C端都各自含有RIPGV和KNFRV序列。并通过系统进化分析,黄梁木AcEXPA家族成员分别属于4个亚家族。
4.筛选到环孢素A受体基因AcCyP作为内参基因用于荧光定量PCR分析。AcEXPA1-16基因在黄梁木中具有明显的组织表达特异性：在四个幼嫩的生长组织中,AcEXPA8在形成层的表达量最高,AcEXPA4、AcEXPA6、AcEXPA15、AcEXPA14和AcEXPA10在嫩叶中大量表达,AcEXPA13只在嫩叶中有所表达,而AcEXPA16仅在嫩根中表达。AcEXPA8在不同生长季节的形成层中的表达也明显不同,7月的最高,1月的最低。在维管发育径向的不同细胞层,AcEXPA8在次生壁形成区的表达量明显低于形成层。
5. AcEXPA8基因以多拷贝的形式在黄梁木基因组中存在。蛋白同源建模显示成熟AcEXPA8蛋白的三维结构与到目前为止仅知的EXP蛋白玉米ZmEXPB1晶体结构极为相似,含有domain1和domain2以及两者之间的连接区域。
6.通过转AcEXPA8基因拟南芥研究,转基因植株与野生型相比,叶片形成加速,叶片面积增大,并且叶片长生长大于宽生长,使得叶片形状变得扁长,在基叶完全成熟期,基座的半径明显增大。而且,转基因植株的高度和茎的粗度也明显大于野生型。
7.过表达AcEXPA8基因促进了拟南芥的次生生长,主要是导致了纤维细胞增长和细胞壁的加厚,但没有改变细胞壁中纤维素的含量。此外,转基因植株茎的髓和皮层薄壁细胞明显增大,而皮层的薄壁细胞在沿周长方向的长度增长尤为明显。
本研究克隆到并分析了黄梁木AcEXPA基因家族成员在不同幼嫩组织的表达差异,对形成层中高丰度表达的AcEXPA8基因的时空表达进行了进一步分析并进行了转基因功能鉴定,结果为今后研究α扩展蛋白基因与团花树木质部生长速度和材质的关系提供基础,并最终为林木分子育种提供潜在的候选基因。
Anthocephalus chinensis with the synonym of Neolamarckia cadamba belonging to the family Rubiaceae, is an evergreen broadleaf arbor and a fast-growing tree species which grows in Tropical Asia and Southern Asia. It was universally accepted as "a miraculous tree" in the World Forestry Congress in1972because of its rapid growth. Meanwhile, it possesses significant development and utilization value in South China. However, genetic information on A. chinensis is still unclear, especially in molecular mechanism study on fast-growing of wood. Expansin is a class of important cell growth regulator, and closely related to the growth and development of plants. So, in the study, we fitstly cloned a-expansin gene sequences in A. chinensis, and inditified the function of the a-expansin gene abundantly expressed in the cambium region, a-expansin genes were isolated by amplifying conservative domain binding with genomic walking and3'RACE techniques from four differential growth tissues in A. chinensis, followed by bioinformatic analysis. Based on these, we selected a suitable reference gene and analysed the temporal and spatial expression pattern of a-expansin genes. AcEXPA8gene expressed highly in cambium region was selected to analyse its copy number in genome, and we also predicted the protein structure and function by homology modeling. Meanwhile, we constructed the plant expression vector and transformed it into Arabidopsis thaliana to further indentify AcEXPA8gene function. The main results and conclusions obtained in this study are shown as follows:
1. AcEXPA1-16genes were cloned by a cloning strategy. Firstly, the conservative regions of A. chinensis a-expansin genes were amplified with the degenerated primers, then, the flank sequences were obtained by genomic walking and3'RACE. Their GenBank accession Nos. were FJ417847, JF922686-JF922700, with corresponding genomic DNA sequences of GenBank accession Nos. GQ228823, JF922701-JF922715.
2. The intron-exon structures of AcEXPAl-16are similar between them, and they all contain intronl and intron2which are conservative in a-expansin genes. All AcEXPAl-16introns conforn with eukaryotic intron splicing pattern of GT-AG, except the intron2of AcEXPA4with TA-GA.
3. All amino acid sequences encoded by AcEXPAI-16gene contain conservative sequences of EXPA protein, including the N-terminal signal peptide, eight C residues, HFD motif and four W residues near the C terminus. But, there is a leucine (L) instead of the first W in AcEXPA12. All mature proteins of AcEXPA1, AcEXPA5, AcEXPA8and AcEXPA9cotain RIPGV sequence at N-terminal and KNFRV sequence at C-terminal. And AcEXPAl-16genes were divided into four subfamilies in the phylogenetic analysis.
4. The screened cyclophilin gene AcCyP was suitable in RT-qPCR analysis as reference gene. AcEXPA1-16genes showed obviously tissue-specific expression in A. chinensis. In the four young tissues in growth, AcEXPA8was expressed highestly in cambium region. AcEXPA4, AcEXPA6, AcEXPA15, AcEXPA14and AcEXPA10were abundantly expressed in the tender leaf. AcEXPA13was only expressed in leaf, whereas AcEXPA16was only expressed in root. The expression of AcEXPA8was significantly different between different growing seasons in cambium region, highest in July, but lowest in January. And the abundance of AcEXPA8expressed in second wall formation region was significantly lower than that in camium region.
5. AcEXPA8gene is present in A. chinensis genomic DNA as multicopy. Protein homology modeling showed that the3D structure of mature protein AcEXPA8is similar to maize ZmEXPB1which is the only known crystal structure of EXP protein so far, and also contains domain1, domain2and connection area between them.
6. Overexpression of AcEXPA8in Arabidopsis could not only accelerate leaf formation, induce leaf area, cause leaf length growth greater than width growth, leading to the leaf shape becoming prolate, and increase the radius of fully mature rosette, but also increase plant height and stem diameter.
7. In transgenic plant stem, overexpression of AcEXPA8promoted secondary growth in Arabidopsis, mainly leading to increase fiber cell length and cell wall thickness, but without changing the content of cellulose in the cell wall. In addition, the area of pith and cortex parenchyma cells increased obviously, and the length of cortex parenchyma cells along the circumference direction increased particularly.
In this study, the AcEXPA gene family member expressions were analyzed in different young tissures, and the temporal and spatial expression pattern of AcEXPA8gene expressed abundantly in cambium region was further analized, followed by gene function analysis. These findings will lay the foundation for studying the relationship between a-expansin genes and growth rate and wood quality of the xylem in A. chinensis, and finally provide potential candidate genes for tree molecular breeding.
引文
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