刺参Apostichopus japonicus (Selenka)对温度胁迫响应分子机理的基础研究
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摘要
以刺参Apostichopus japonicus为研究对象,探讨了温度胁迫下刺参基因表达变化,构建了高温刺激与未经温度刺激的刺参体壁消减cDNA文库,克隆了与温度密切相关的热休克蛋白90和热休克蛋白26基因cDNA全长序列,分析了基因结构,在mRNA水平上研究了两个基因在不同刺激温度、刺激时间及不同组织中的表达情况,并评价了不同群体刺参耐温性状的差异。
1.构建了温度胁迫条件下刺参体壁消减cDNA文库。以温度刺激条件下刺参体壁为检测子(tester),以对照组刺参体壁为驱动子(driver),经两轮差减杂交,两轮抑制性PCR扩增,获得正向消减文库;以温度刺激条件下刺参体壁为驱动子(driver),以对照组刺参体壁为检测子(tester),构建反向消减文库。在正反向文库中随机挑选768个克隆进行PCR扩增鉴定,发现多数克隆都含有插入片段,进一步利用斑点杂交进行筛选,得到737个有差异表达的阳性克隆,经测序共获得165个EST序列,其中65个为已知基因,100个为未知基因。对这些基因进行功能分类,发现正向库中免疫应激相关基因数量明显多于反向库,而能量代谢相关基因明显少于反向库,表明温度胁迫诱导应激防御相关基因的表达,而抑制代谢相关基因的转录。
2.克隆了刺参热休克蛋白90(Aphsp90)和热休克蛋白26(Aphsp26)cDNA序列全长。刺参热休克蛋白90(Aphsp90)和热休克蛋白26(Aphsp26)cDNA全长分别为3458bp和1688bp,编码720和236个氨基酸,均为不具有信号肽的胞内蛋白。多重序列比对、系统进化分析显示ApHsp90和ApHsp26氨基酸序列与其他物种Hsp90和sHSPs具有同源性,蛋白质三维结构研究发现这两个氨基酸序列具有Hsp90和sHSPs的特征功能区域,表明获得的cDNA序列为目的基因。
3.揭示了热休克蛋白90(Aphsp90)和热休克蛋白26(Aphsp26)基因表达特征。应用RT-PCR检测Aphsp90和Aphsp26基因在不同温度下(20、22、24、26℃)刺参不同组织的表达发现,刺参体壁、呼吸树和肠道中Aphsp90和Aphsp26基因表达发生变化以响应温度胁迫,且响应具有组织差异性。在20℃对照组中,不同组织中基因转录水平高低顺序为刺参肠道>呼吸树>体壁;随着温度的升高,Aphsp90和Aphsp26基因表达量与温度呈现正相关性,在不同组织中都显著升高,肠道中两个基因转录水平变化幅度最大,肠道可能是刺参机体热敏感性较高的组织。基因时序表达结果显示两个基因转录水平与胁迫时间具有较好的正相关性,Aphsp26基因转录水平在胁迫初期(0-6h)显著升高,之后下降,末期恢复到接近对照水平,Aphsp90基因表达量在胁迫4h达到最大值,6h维持在最大值,之后缓慢下降恢复到接近对照水平。
4.比较了高温定向选育后南移养殖的浙江群体和烟台野生群体对高温的耐受性。高温(28℃)培育对不同群体幼体存活率、变态率及特定生长率的影响结果显示,在高温条件下两个群体幼体存活率、变态率及特定生长率都比恒温条件(22℃)显著下降,浙江南移群体在高温条件下表现出较好的耐受性,各类指标都高于烟台野生群体。分析了两个群体幼体温度胁迫下的生长趋势,发现随着温度的升高,幼体发育加快,提前进入变态期,浙江幼体在高温条件下最大体长增长,而烟台幼体最大体长缩短。对比不同群体幼参32℃热激17天后的存活率,发现浙江南移群体比烟台野生群体表现出较高的存活率;分析不同群体Aphsp90和Aphsp26基因在28℃热激后的表达情况,结果显示浙江南移群体两个基因转录水平在热激后各个时间点都低于烟台群体,表明刺参耐受性差异与Aphsp90和Aphsp26基因表达模式存在一定的相关性。
In order to understand the response to heat stress at the molecular level in Apostichopus japonicus, a suppression subtractive hybridization library was constructed to analyze the difference expression in the body wall of A. japonicus. The full-length cDNAs of Aphsp90 and Aphsp26 were cloned and analyzed, and the expression profiles of Aphsp90 and Aphsp26 in different tissues under different temperatures and length of times were determined by real time quantitative PCR. Further the heat tolerance of different populations was assessed.
1. Suppression subtractive hybridization library was constructed to analyze the differences in gene expression in the body wall of A. japonicus induced by heat stress (28°C). The forward SSH library used the cDNA from the heat-treated group as the tester and the cDNA from the control as the driver in order to find the up-regulated genes. The reverse SSH library was used to identify the down-regulated genes, using the cDNA from the control as a tester and the cDNA from the heat-treated group as a driver.The 737 clones were proved as positive clones with bacterial colony PCR and dot hybridization, and 165 distinct sequences were obtained, with 65 sharing similarity to sequences in public databases, 100 sharing no similarity to sequences in public databases. The sequenced genes were classified into different functional categories, and it was found that increased expression of some genes in response to stress was concomitant with decreased expression of genes concerned with energy production and metabolism under heat stress in A. japonicus. It indicates that heat stress could induce the expression of some genes with the function of immunity but suppress the expression of gene concerned with metabolism.
2. The genes of Aphsp90 and Aphsp26 were cloned. The full-length cDNA of Aphsp90 and Aphsp26 contains 3,458bp and 1,688bp encoding 720 and 236 amino acids separately. Multiple alignments and phylogenetic analysis indicated that the animo acid sequences of ApHsp90 and ApHsp26 shared high similarity with Hsp90 and sHSPs sequences of other species. The three-dimensional structure of ApHsp90 contains two characteristic domains of Hsp90 which indicated the same function of ApHsp90, the structure analysis of ApHsp26 showed thatα-crystallin domain which is the most conserved elements in sHSPs is presented in this sequence. It indicates the validity of these sequences.
3. The expression characteristics of Aphsp90 and Aphsp26 were analyzed. The expression patterns of these two genes in different tissues of A. japonicus exposure in four temperature treatments (20, 22, 24, 26°C) were determined by real time quantitative PCR. It was found that the messenger RNA (mRNA) transcripts of these two genes varied among different tissues. Under normal condition and heat shock, the mRNA expression of these two genes was higher in intestine compared to other tissues. Heat shock could accelerate the gene expression of Aphsp90 and Aphsp26, with the increase of temperature the expression of Aphsp90 and Aphsp26 mRNA elevated, especially in the intestine of A. japonicus. It indicates that the intestine of A. japonicus is the heat-sensitive tissue.The relationship between expression level and duration of heat shock was determined, and the results showed that the level of Aphsp90 and Aphsp26 was time-dependent. The transcripts of Aphsp26 increased significantly in the first few hours, reached maximum expression after heat shock for 6h then decreased gradually and reached the pretreatment level at 48h. The mRNA level of Aphsp90 reached a peak at 4 h and remained high at 6 h under heat stress, then decreased gradually to the pretreatment level at 48 h.
4. The heat tolerance of different populations of A. japonicus was analyzed. Yantai population and Zhejiang population experiencing high-temperature breeding and south breeding were chosen as research objects. The survival rate, metamorphosis rate and special growth rate of larvae in these two populations under heat stress were determined. The results showed that heat shock could influence the survival of larvae, these indexes in heat group were lower than in control group in both populations, but the larvae of Zhejiang population exhibited higher heat tolerance compared to Yantai population with higher survival rate, metamorphosis rate and special growth rate. Under heat shock the development of larvae increased, with advance into the doliolarias stage. The maximum length of larvae in Yantai population shortened under heat shock, but the variation trend was opposite in Zhejiang population. Juveniles of both populations were maintained at 32°C for 17 days to determine the survival rate. It was indicated that Zhejiang population exhibited higher survival rate than Yantai population under heat shock. Further in molecular level the difference in gene expression of Aphsp90 and Aphsp26 in these two populations under 28°C heat shock was studied. The results indicated that the transcripts of Aphsp90 and Aphsp26 mRNA in Zhejiang population were always lower than Yantai population at different time of heat shock. It indicates that the difference in heat tolerance has relationship with the expression profiles of Aphsp90 and Aphsp26 mRNA.
1.构建了温度胁迫条件下刺参体壁消减cDNA文库。以温度刺激条件下刺参体壁为检测子(tester),以对照组刺参体壁为驱动子(driver),经两轮差减杂交,两轮抑制性PCR扩增,获得正向消减文库;以温度刺激条件下刺参体壁为驱动子(driver),以对照组刺参体壁为检测子(tester),构建反向消减文库。在正反向文库中随机挑选768个克隆进行PCR扩增鉴定,发现多数克隆都含有插入片段,进一步利用斑点杂交进行筛选,得到737个有差异表达的阳性克隆,经测序共获得165个EST序列,其中65个为已知基因,100个为未知基因。对这些基因进行功能分类,发现正向库中免疫应激相关基因数量明显多于反向库,而能量代谢相关基因明显少于反向库,表明温度胁迫诱导应激防御相关基因的表达,而抑制代谢相关基因的转录。
2.克隆了刺参热休克蛋白90(Aphsp90)和热休克蛋白26(Aphsp26)cDNA序列全长。刺参热休克蛋白90(Aphsp90)和热休克蛋白26(Aphsp26)cDNA全长分别为3458bp和1688bp,编码720和236个氨基酸,均为不具有信号肽的胞内蛋白。多重序列比对、系统进化分析显示ApHsp90和ApHsp26氨基酸序列与其他物种Hsp90和sHSPs具有同源性,蛋白质三维结构研究发现这两个氨基酸序列具有Hsp90和sHSPs的特征功能区域,表明获得的cDNA序列为目的基因。
3.揭示了热休克蛋白90(Aphsp90)和热休克蛋白26(Aphsp26)基因表达特征。应用RT-PCR检测Aphsp90和Aphsp26基因在不同温度下(20、22、24、26℃)刺参不同组织的表达发现,刺参体壁、呼吸树和肠道中Aphsp90和Aphsp26基因表达发生变化以响应温度胁迫,且响应具有组织差异性。在20℃对照组中,不同组织中基因转录水平高低顺序为刺参肠道>呼吸树>体壁;随着温度的升高,Aphsp90和Aphsp26基因表达量与温度呈现正相关性,在不同组织中都显著升高,肠道中两个基因转录水平变化幅度最大,肠道可能是刺参机体热敏感性较高的组织。基因时序表达结果显示两个基因转录水平与胁迫时间具有较好的正相关性,Aphsp26基因转录水平在胁迫初期(0-6h)显著升高,之后下降,末期恢复到接近对照水平,Aphsp90基因表达量在胁迫4h达到最大值,6h维持在最大值,之后缓慢下降恢复到接近对照水平。
4.比较了高温定向选育后南移养殖的浙江群体和烟台野生群体对高温的耐受性。高温(28℃)培育对不同群体幼体存活率、变态率及特定生长率的影响结果显示,在高温条件下两个群体幼体存活率、变态率及特定生长率都比恒温条件(22℃)显著下降,浙江南移群体在高温条件下表现出较好的耐受性,各类指标都高于烟台野生群体。分析了两个群体幼体温度胁迫下的生长趋势,发现随着温度的升高,幼体发育加快,提前进入变态期,浙江幼体在高温条件下最大体长增长,而烟台幼体最大体长缩短。对比不同群体幼参32℃热激17天后的存活率,发现浙江南移群体比烟台野生群体表现出较高的存活率;分析不同群体Aphsp90和Aphsp26基因在28℃热激后的表达情况,结果显示浙江南移群体两个基因转录水平在热激后各个时间点都低于烟台群体,表明刺参耐受性差异与Aphsp90和Aphsp26基因表达模式存在一定的相关性。
In order to understand the response to heat stress at the molecular level in Apostichopus japonicus, a suppression subtractive hybridization library was constructed to analyze the difference expression in the body wall of A. japonicus. The full-length cDNAs of Aphsp90 and Aphsp26 were cloned and analyzed, and the expression profiles of Aphsp90 and Aphsp26 in different tissues under different temperatures and length of times were determined by real time quantitative PCR. Further the heat tolerance of different populations was assessed.
1. Suppression subtractive hybridization library was constructed to analyze the differences in gene expression in the body wall of A. japonicus induced by heat stress (28°C). The forward SSH library used the cDNA from the heat-treated group as the tester and the cDNA from the control as the driver in order to find the up-regulated genes. The reverse SSH library was used to identify the down-regulated genes, using the cDNA from the control as a tester and the cDNA from the heat-treated group as a driver.The 737 clones were proved as positive clones with bacterial colony PCR and dot hybridization, and 165 distinct sequences were obtained, with 65 sharing similarity to sequences in public databases, 100 sharing no similarity to sequences in public databases. The sequenced genes were classified into different functional categories, and it was found that increased expression of some genes in response to stress was concomitant with decreased expression of genes concerned with energy production and metabolism under heat stress in A. japonicus. It indicates that heat stress could induce the expression of some genes with the function of immunity but suppress the expression of gene concerned with metabolism.
2. The genes of Aphsp90 and Aphsp26 were cloned. The full-length cDNA of Aphsp90 and Aphsp26 contains 3,458bp and 1,688bp encoding 720 and 236 amino acids separately. Multiple alignments and phylogenetic analysis indicated that the animo acid sequences of ApHsp90 and ApHsp26 shared high similarity with Hsp90 and sHSPs sequences of other species. The three-dimensional structure of ApHsp90 contains two characteristic domains of Hsp90 which indicated the same function of ApHsp90, the structure analysis of ApHsp26 showed thatα-crystallin domain which is the most conserved elements in sHSPs is presented in this sequence. It indicates the validity of these sequences.
3. The expression characteristics of Aphsp90 and Aphsp26 were analyzed. The expression patterns of these two genes in different tissues of A. japonicus exposure in four temperature treatments (20, 22, 24, 26°C) were determined by real time quantitative PCR. It was found that the messenger RNA (mRNA) transcripts of these two genes varied among different tissues. Under normal condition and heat shock, the mRNA expression of these two genes was higher in intestine compared to other tissues. Heat shock could accelerate the gene expression of Aphsp90 and Aphsp26, with the increase of temperature the expression of Aphsp90 and Aphsp26 mRNA elevated, especially in the intestine of A. japonicus. It indicates that the intestine of A. japonicus is the heat-sensitive tissue.The relationship between expression level and duration of heat shock was determined, and the results showed that the level of Aphsp90 and Aphsp26 was time-dependent. The transcripts of Aphsp26 increased significantly in the first few hours, reached maximum expression after heat shock for 6h then decreased gradually and reached the pretreatment level at 48h. The mRNA level of Aphsp90 reached a peak at 4 h and remained high at 6 h under heat stress, then decreased gradually to the pretreatment level at 48 h.
4. The heat tolerance of different populations of A. japonicus was analyzed. Yantai population and Zhejiang population experiencing high-temperature breeding and south breeding were chosen as research objects. The survival rate, metamorphosis rate and special growth rate of larvae in these two populations under heat stress were determined. The results showed that heat shock could influence the survival of larvae, these indexes in heat group were lower than in control group in both populations, but the larvae of Zhejiang population exhibited higher heat tolerance compared to Yantai population with higher survival rate, metamorphosis rate and special growth rate. Under heat shock the development of larvae increased, with advance into the doliolarias stage. The maximum length of larvae in Yantai population shortened under heat shock, but the variation trend was opposite in Zhejiang population. Juveniles of both populations were maintained at 32°C for 17 days to determine the survival rate. It was indicated that Zhejiang population exhibited higher survival rate than Yantai population under heat shock. Further in molecular level the difference in gene expression of Aphsp90 and Aphsp26 in these two populations under 28°C heat shock was studied. The results indicated that the transcripts of Aphsp90 and Aphsp26 mRNA in Zhejiang population were always lower than Yantai population at different time of heat shock. It indicates that the difference in heat tolerance has relationship with the expression profiles of Aphsp90 and Aphsp26 mRNA.
引文
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