日本血吸虫性别差异表达基因的筛选及新基因的克隆分析
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摘要
血吸虫病是由血吸虫感染引起的一种分布广泛、危害严重的人畜共患寄生虫病。血吸虫为吸虫中罕见的雌雄异体,性别的分化对血吸虫的发育及致病起着至关重要的作用。雌雄虫合抱是血吸虫雌虫发育成熟和产卵的关键,成熟雌虫所产生大量虫卵而引起肝脏病变是造成宿主主要病理损害和疾病再传播的前提。基于此本论文开展了日本血吸虫性别差异表达基因的筛选研究,并对部分差异表达新基因进行了克隆分析,以探索血吸虫性别发育的分子机理,开拓免疫预防新途径。
应用抑制性消减杂交技术构建了日本血吸虫雄虫和雌虫的消减杂交文库。消减效率分析表明同源基因经过消减杂交后得到了有效扣除,消减效率较高;重组比率及插入片断长度分析结果表明雌虫消减文库的重组比率为92%,且85%的插入片断大于500bp;雄虫消减文库的重组比率为91%,且90%以上的插入片断大于500bp;对构建的日本血吸虫雌雄虫消减库随机挑选部分克隆测序,分别获得64条雌虫EST序列和62条雄虫EST序列。同源性搜索结果表明分别代表了45和48个日本血吸虫基因,雌虫和雄虫消减文库的冗余率分别为29.7%和22.6%;所测EST序列共有25条与12种血吸虫已知基因匹配,其中雌虫消减文库中与血吸虫已知基因高度同源的有日本血吸虫卵壳蛋白、卵黄铁蛋白Ferrtin-1、fs800等基因。雄虫消减文库与已知血吸虫基因高度同源的有肌动蛋白、原肌球蛋白等基因。这几种血吸虫基因已经被证实为性别差异表达基因,结果表明本研究所构建的日本血吸虫性别差异消减文库质量较高,可以用于性别差异表达基因的筛选研究。
利用cDNA微阵列技术在整个基因组水平上对日本血吸虫性别差异表达基因进行筛选。从构建好的雌虫和雄虫两个消减文库中共挑选3072个克隆制成cDNA微阵列。芯片杂交后获得953个雌虫高表达和1014个雄虫高表达的基因克隆。其中855个克隆经测序分析代表297个单基因,雌雄虫分别为137个和160个。序列同源性搜索结果表明:297个单基因中,247个与已知EST有95%～100%同源性;雌虫所代表的137个单基因中,蛋白功能已知的血吸虫基因有18个,蛋白功能未知的血吸虫基因96个,与已知EST高度同源的基因13个,其它物种基因2个,没有同源性的新EST 8个。雄虫所代表的160个血吸虫单基因中,蛋白功能已知的血吸虫基因30个,蛋白功能未知的血吸虫基因109个,与已知EST高度同源的基因15个,其它物种基因1个,没有同源性的新EST 5个。从中选取7个非冗余的差异表达基因克隆,用实时定量PCR进行了验证,证明上述克隆确为血吸虫性别差异表达的基因克隆,与芯片杂交结果一致。所测EST编码蛋白的功能预测结果显示:雌虫消减文库中与已知血吸虫基因高度同源的有卵壳蛋白、卵黄铁蛋白、fs800、RXR受体、组织蛋白酶B和组织蛋白酶L、过氧化物歧化酶(SOD)、谷胱甘肽还原酶等,提示多与血吸虫的产卵、能量代谢、转录调节及抗氧化反应相关。雄虫消减文库与已知血吸虫基因高度同源的有肌动蛋白、原肌球蛋白、肌球蛋白、胶原、22.6ku膜蛋白、23ku膜蛋白、钙蛋白酶、calponin、钙调蛋白、钙网蛋白、热激蛋白HSP60、HSP70、HSP90等。EST的编码蛋白功能预测结果提示这类基因多为虫体的结构蛋白与虫体运动相关蛋白,参与离子调节与信号转导、蛋白质的合成、转运及分解代谢等。
应用RACE技术扩增了4个血吸虫全长新基因cDNA。并利用生物信息学分析工具对新基因编码蛋白的理化性质、跨膜结构、抗原位点进行了分析预测,对编码蛋白的保守功能域、结构位点及其同源性蛋白进行了搜索。结果显示雌虫高表达f2基因编码蛋白具有泛素和核糖体蛋白S30e的保守功能域。雄虫高表达cm3基因编码蛋白具有C2H2型锌指结构的保守功能域,cm4编码蛋白具有SPla/RYanodine receptor SPRY结构域。m1基因编码蛋白可能是一种血吸虫特有蛋白。对新基因生物学功能的深入研究,对于探索血吸虫性别发育的分子机制、开发高效候选疫苗和药物新靶标具有重要价值。
Schistosomiasis caused by Schistosoma japonicum is an important zoonosis in China and southeast Asia. A virtually unique trait of all species of Schistosoma (phylum Platyhelminthes) is their evolution from hermaphrodite ancestors into sexually dimorphic species. Germ-cell differentiation is crucial to the developments of schistosome. Pairing between male and female worm of schistosome is necessary for female reproductive development and for the maintenance of her mature state. Adult female worm can produce a large number of eggs which cause the main pathological changes in the infected host and also are responsible for the dissemination of parasite. Based on above characteristics of schistosome, the screening and identification of gender-associated differentially expressed genes of S. japonicum were carried out, and several differentially expressed new genes were cloned and analyzed to probe the molecular mechanism of S. japonicum gender-associated gene development that may be considered new targets for vaccination.
1. Construction and analysis of suppression subtractive hybridization libraries of S. japanicum female and male worms
In this study, Poly A+RNA was isolated respectively from S. japonicum (Sj) female and male adult worms, and suppression subtractive hybridization(SSH) libraries of Sj female and male worms were constructed by using Clontech PCR-selectTM cDNA subtraction kit. The evaluation results indicated that the two SSH libraries were high quality, and were applicable to isolate Sj differentially expressed genes on the whole genome scale. The recombinant rates of female and male SSH libraries were 92% and 91% respectively. Some clones were selected randomly and sequenced, 64 female ESTs and 62 male ESTs were obtained. The nonredundancy rate of the female and male SSH libraries were 29.7% and 22.6% respectively. The homology search results suggest that there were 45 female and 48 male unigenes .There were egg shell protein ,ferrtin-1and female-specific 800 in female SSH library and actin, trypomysin in male SSH library. These genes were previously verified as gender-associated differentially expressed genes by other researchers. Identifying the more differentially expressed genes may uncover molecular mechanism of female and male sexual maturation, It was an important step in developing strategies for blocking the worm’s reproductive life cycle, and their potential to induce host morbidity and ultimately preventing successful transmission.
2. Analysis of the gender-associated gene differentially expression profile of S. japonicum using cDNA microarray technique
Four pieces of S. japonicum gender-associated differentially expression cDNA microarrays were fabricated. Each cDNA microarray contained 1536 female and 1536 male cDNA clones originating from SSH libraries. Microarray hybridization results suggested that there were 953 female and 1014 male differentially expression gene clones.855 gene clones were selected and sequenced. The results of sequence analysis indicated that these gene clones represented 297unigenes composed of 137 female and 160 male unigenes. 247 genes were homologous to known ESTs in GenBank by the degree from 95% to 100%. Among 137 female unigenes,18 genes had known protein functions ,96 genes had unknown protein functions, 13 genes had known EST match, 8 genes represented unknown new genes of Schistosoma which had no database match. Among 160 male unigenes,30 genes had known protein functions, 109 genes had unknown protein functions, 15 genes had known EST match, 5 genes represented unknown new genes of Schistosoma which had no database match. Seven nonredundant differential expression gene clones were verified by Real-time PCR, which demonstrated that these clones were indeed differentially expressed, and confirmed the cDNA microarray results, thereby supporting the reliability of the system. Our results indicated that it is high efficient to isolate and identify differentially expressed genes by using SSH combined with cDNA microarray.
Homology of sequenced ESTs were analyzed and functions of genes encoding protein were predicted. The results indicated that there were egg shell protein, ferrtin-1, RXR receptor, cathepsin B, cathepsin L, SOD, Glutathione reductase in female SSH library, and there were actin, tropomysin, myosin, collagen, Sj 22.6 kDa membrane- associated antigen, Sj 23 kDa membrane-associated protein, calpain、calponin、calmodulin、calreticulin、HSP60、HSP70、HSP90 etc. in male SSH library. The results suggested that genes from female SSH library mainly involved in the process of egg production, metabolism, transcription regulation and anti-oxidation, The genes from male SSH library were intimately involved in the structural organization including components of the muscular system, tegumental proteins and the underlying cytoskeleton, ion regulation, signal transduction, protein biosynthesis and catabolism. This apparent division of labour between the genders had an extensive transcriptional basis. The male ensured the survival of the egg-laying female by providing physical support and musculature to aid feeding, physical transportation within the vasculature. The female then concentrated energy expenditure on egg production.
3. Cloning and analysis of gender-associated differentially expressed new genes of S. japonicum
Full-length cDNA of the four new gender-associated differentially expressed genes were obtained by the RACE (rapid amplification of cDNA ends) technique. F2 was up-regulated in female worms, and cm3, cm4, m1 were up-regulated in male worms. With bioinformatics tools, protein encoded by these new genes were predicted in these, including their physical and chemical property, trans-membrane structure, antigen site, conserved domain, function site and homologous protein search. The results indicated that f2 contained conserved domains of ubiquitin and ribosomal protein S30 which may involve in protein biosynthesis and modification. Cm3 contained conserved domains of C2H2-type zinc-finger which may involve in nucleic acid binding, cm4 contained conserved domains of SPla/RYanodine receptor SPRY which mediate Ca2+-release from the sarcoplasmic or endoplasmic reticulum and the intracellular Ca2+ store.
Investigation on biological function of these new genes will help to better understand the molecular mechanisms regulating schistosome sex developmental processes, and to establish a basis for exploiting effective vaccine candidate and new drug target.
应用抑制性消减杂交技术构建了日本血吸虫雄虫和雌虫的消减杂交文库。消减效率分析表明同源基因经过消减杂交后得到了有效扣除,消减效率较高;重组比率及插入片断长度分析结果表明雌虫消减文库的重组比率为92%,且85%的插入片断大于500bp;雄虫消减文库的重组比率为91%,且90%以上的插入片断大于500bp;对构建的日本血吸虫雌雄虫消减库随机挑选部分克隆测序,分别获得64条雌虫EST序列和62条雄虫EST序列。同源性搜索结果表明分别代表了45和48个日本血吸虫基因,雌虫和雄虫消减文库的冗余率分别为29.7%和22.6%;所测EST序列共有25条与12种血吸虫已知基因匹配,其中雌虫消减文库中与血吸虫已知基因高度同源的有日本血吸虫卵壳蛋白、卵黄铁蛋白Ferrtin-1、fs800等基因。雄虫消减文库与已知血吸虫基因高度同源的有肌动蛋白、原肌球蛋白等基因。这几种血吸虫基因已经被证实为性别差异表达基因,结果表明本研究所构建的日本血吸虫性别差异消减文库质量较高,可以用于性别差异表达基因的筛选研究。
利用cDNA微阵列技术在整个基因组水平上对日本血吸虫性别差异表达基因进行筛选。从构建好的雌虫和雄虫两个消减文库中共挑选3072个克隆制成cDNA微阵列。芯片杂交后获得953个雌虫高表达和1014个雄虫高表达的基因克隆。其中855个克隆经测序分析代表297个单基因,雌雄虫分别为137个和160个。序列同源性搜索结果表明:297个单基因中,247个与已知EST有95%～100%同源性;雌虫所代表的137个单基因中,蛋白功能已知的血吸虫基因有18个,蛋白功能未知的血吸虫基因96个,与已知EST高度同源的基因13个,其它物种基因2个,没有同源性的新EST 8个。雄虫所代表的160个血吸虫单基因中,蛋白功能已知的血吸虫基因30个,蛋白功能未知的血吸虫基因109个,与已知EST高度同源的基因15个,其它物种基因1个,没有同源性的新EST 5个。从中选取7个非冗余的差异表达基因克隆,用实时定量PCR进行了验证,证明上述克隆确为血吸虫性别差异表达的基因克隆,与芯片杂交结果一致。所测EST编码蛋白的功能预测结果显示:雌虫消减文库中与已知血吸虫基因高度同源的有卵壳蛋白、卵黄铁蛋白、fs800、RXR受体、组织蛋白酶B和组织蛋白酶L、过氧化物歧化酶(SOD)、谷胱甘肽还原酶等,提示多与血吸虫的产卵、能量代谢、转录调节及抗氧化反应相关。雄虫消减文库与已知血吸虫基因高度同源的有肌动蛋白、原肌球蛋白、肌球蛋白、胶原、22.6ku膜蛋白、23ku膜蛋白、钙蛋白酶、calponin、钙调蛋白、钙网蛋白、热激蛋白HSP60、HSP70、HSP90等。EST的编码蛋白功能预测结果提示这类基因多为虫体的结构蛋白与虫体运动相关蛋白,参与离子调节与信号转导、蛋白质的合成、转运及分解代谢等。
应用RACE技术扩增了4个血吸虫全长新基因cDNA。并利用生物信息学分析工具对新基因编码蛋白的理化性质、跨膜结构、抗原位点进行了分析预测,对编码蛋白的保守功能域、结构位点及其同源性蛋白进行了搜索。结果显示雌虫高表达f2基因编码蛋白具有泛素和核糖体蛋白S30e的保守功能域。雄虫高表达cm3基因编码蛋白具有C2H2型锌指结构的保守功能域,cm4编码蛋白具有SPla/RYanodine receptor SPRY结构域。m1基因编码蛋白可能是一种血吸虫特有蛋白。对新基因生物学功能的深入研究,对于探索血吸虫性别发育的分子机制、开发高效候选疫苗和药物新靶标具有重要价值。
Schistosomiasis caused by Schistosoma japonicum is an important zoonosis in China and southeast Asia. A virtually unique trait of all species of Schistosoma (phylum Platyhelminthes) is their evolution from hermaphrodite ancestors into sexually dimorphic species. Germ-cell differentiation is crucial to the developments of schistosome. Pairing between male and female worm of schistosome is necessary for female reproductive development and for the maintenance of her mature state. Adult female worm can produce a large number of eggs which cause the main pathological changes in the infected host and also are responsible for the dissemination of parasite. Based on above characteristics of schistosome, the screening and identification of gender-associated differentially expressed genes of S. japonicum were carried out, and several differentially expressed new genes were cloned and analyzed to probe the molecular mechanism of S. japonicum gender-associated gene development that may be considered new targets for vaccination.
1. Construction and analysis of suppression subtractive hybridization libraries of S. japanicum female and male worms
In this study, Poly A+RNA was isolated respectively from S. japonicum (Sj) female and male adult worms, and suppression subtractive hybridization(SSH) libraries of Sj female and male worms were constructed by using Clontech PCR-selectTM cDNA subtraction kit. The evaluation results indicated that the two SSH libraries were high quality, and were applicable to isolate Sj differentially expressed genes on the whole genome scale. The recombinant rates of female and male SSH libraries were 92% and 91% respectively. Some clones were selected randomly and sequenced, 64 female ESTs and 62 male ESTs were obtained. The nonredundancy rate of the female and male SSH libraries were 29.7% and 22.6% respectively. The homology search results suggest that there were 45 female and 48 male unigenes .There were egg shell protein ,ferrtin-1and female-specific 800 in female SSH library and actin, trypomysin in male SSH library. These genes were previously verified as gender-associated differentially expressed genes by other researchers. Identifying the more differentially expressed genes may uncover molecular mechanism of female and male sexual maturation, It was an important step in developing strategies for blocking the worm’s reproductive life cycle, and their potential to induce host morbidity and ultimately preventing successful transmission.
2. Analysis of the gender-associated gene differentially expression profile of S. japonicum using cDNA microarray technique
Four pieces of S. japonicum gender-associated differentially expression cDNA microarrays were fabricated. Each cDNA microarray contained 1536 female and 1536 male cDNA clones originating from SSH libraries. Microarray hybridization results suggested that there were 953 female and 1014 male differentially expression gene clones.855 gene clones were selected and sequenced. The results of sequence analysis indicated that these gene clones represented 297unigenes composed of 137 female and 160 male unigenes. 247 genes were homologous to known ESTs in GenBank by the degree from 95% to 100%. Among 137 female unigenes,18 genes had known protein functions ,96 genes had unknown protein functions, 13 genes had known EST match, 8 genes represented unknown new genes of Schistosoma which had no database match. Among 160 male unigenes,30 genes had known protein functions, 109 genes had unknown protein functions, 15 genes had known EST match, 5 genes represented unknown new genes of Schistosoma which had no database match. Seven nonredundant differential expression gene clones were verified by Real-time PCR, which demonstrated that these clones were indeed differentially expressed, and confirmed the cDNA microarray results, thereby supporting the reliability of the system. Our results indicated that it is high efficient to isolate and identify differentially expressed genes by using SSH combined with cDNA microarray.
Homology of sequenced ESTs were analyzed and functions of genes encoding protein were predicted. The results indicated that there were egg shell protein, ferrtin-1, RXR receptor, cathepsin B, cathepsin L, SOD, Glutathione reductase in female SSH library, and there were actin, tropomysin, myosin, collagen, Sj 22.6 kDa membrane- associated antigen, Sj 23 kDa membrane-associated protein, calpain、calponin、calmodulin、calreticulin、HSP60、HSP70、HSP90 etc. in male SSH library. The results suggested that genes from female SSH library mainly involved in the process of egg production, metabolism, transcription regulation and anti-oxidation, The genes from male SSH library were intimately involved in the structural organization including components of the muscular system, tegumental proteins and the underlying cytoskeleton, ion regulation, signal transduction, protein biosynthesis and catabolism. This apparent division of labour between the genders had an extensive transcriptional basis. The male ensured the survival of the egg-laying female by providing physical support and musculature to aid feeding, physical transportation within the vasculature. The female then concentrated energy expenditure on egg production.
3. Cloning and analysis of gender-associated differentially expressed new genes of S. japonicum
Full-length cDNA of the four new gender-associated differentially expressed genes were obtained by the RACE (rapid amplification of cDNA ends) technique. F2 was up-regulated in female worms, and cm3, cm4, m1 were up-regulated in male worms. With bioinformatics tools, protein encoded by these new genes were predicted in these, including their physical and chemical property, trans-membrane structure, antigen site, conserved domain, function site and homologous protein search. The results indicated that f2 contained conserved domains of ubiquitin and ribosomal protein S30 which may involve in protein biosynthesis and modification. Cm3 contained conserved domains of C2H2-type zinc-finger which may involve in nucleic acid binding, cm4 contained conserved domains of SPla/RYanodine receptor SPRY which mediate Ca2+-release from the sarcoplasmic or endoplasmic reticulum and the intracellular Ca2+ store.
Investigation on biological function of these new genes will help to better understand the molecular mechanisms regulating schistosome sex developmental processes, and to establish a basis for exploiting effective vaccine candidate and new drug target.
引文
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