鸭卵泡发育差异表达基因抑制性消减文库的构建及分析
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摘要
禽类的产蛋率,除了其他方面外,取决于等级前卵泡能被有效地募集进入等级卵泡,即通过卵泡的生长和闭锁建立卵泡等级。随着日龄的老化产蛋率的下降归因于卵泡闭锁的增加以及可供选择进入最后生长阶段(最大等级卵泡／排卵前卵泡)的小/生长卵泡数量减少。对人和哺乳动物巢卵泡发育研究表明,卵泡的生长和分化是多因素调控的复杂的生理生化过程,不仅受生殖轴内分泌激素的调控,而且卵巢中许多局部的生长、分化因子也以自分泌或旁分泌等形式发挥重要作用。目前,对于禽类卵泡发育的研究,虽然已经知道一些基因在不同发育阶段的卵泡中差异表达,但是对于复杂生命体而言,仅仅是其中很小部分,仍然没有合理假说来说明禽类卵泡等级的建立机制。通过筛选鸭卵泡差异表达的基因,比较不同发育阶段卵泡的差异表达基因,可以对这些差异基因在卵泡发育和分化过程中所发挥的作用有更深入的了解。揭示卵泡在不同生理状况下差异基因的表达情况,为进一步研究禽类性成熟、产蛋的启动、产蛋的维持和高产性能的分子生物学机制提供重要资料,并为最终揭示禽类卵泡发育和等级建立的分子机制,从而在基因水平上进行遗传操作,延长产蛋高峰期,提高鸭的产蛋率提供理论依据。
本实验以产蛋高峰期绍兴鸭母鸭不同阶段的卵泡为实验材料,采用DSN酶介导的均一化消减杂交方法构建鸭卵泡差异表达基因的消减cDNA文库,以期找到与鸭卵泡发育相关的基因,探索卵泡等级建立与分化的基因表达调控的分子机制。本实验构建了两个鸭卵泡差异表达基因的双向消减文库：
以最大等级卵泡和等级前卵泡这两个不同发育阶段的卵泡为实验材料进行消减杂交。其中,以最大等级卵泡为Tester,等级前卵泡为Driver,建立的为正向消减文库；以等级前卵泡为Tester最大等级卵泡为Driver建立的为反向消减文库。实验过程中,总RNA与反转录成的cDNA质量较好,消减产物得到很好的富集。正向消减文库中获得了88条独立的差异表达的基因,BLAST比对结果显示,其中60条具有同源基因,28条相似度极低或无同源性的基因。筛选出与卵泡发育相关基因LHR和EGFR,与细胞增殖和分化有关的基因SLC家族等,影响胚胎发育的NADH1α脱氢酶同源基因NADH1β几个编码激酶或其同源基因如PRKG2、 AKAP2、similar to protein kinase D1。此外,筛选到我们还分离了一些未知功能的基因。反向消减文库中获得了72条独立的差异表达的基因,BLAST比对结果显示,其中51条具有同源基因,21条无相似或者同源性极低的基因。筛选出的基因有RBMII,拼接因子SF2亚单位p32和U2AF35kDa subunit-like, tensin基因,真核生物翻译起始基因EIF3I,微管组分基因α-tubu1in,肠平滑肌肌动蛋白gamma2,含靶蛋白同源基因similar to palladin,免疫反应相关基因CD58,编码微小染色体维持蛋白的基因MCM5, RCC2,参与细胞侵袭和转移的基因Flotillin-2,酶类(包括激酶)及其同源基因AFG3ATPase f ami ly gene3-like2、AK3L2、TAOK3等基因。另外,我们还分离了一些未知功能的新基因。
以最大等级卵泡和最小等级卵泡这两个不同发育阶段的卵泡为实验材料进行消减杂交。其中,以最大等级卵泡为Tester,最小等级卵泡为Driver,建立的为正向消减文库；以最小等级卵泡为Tester,最大等级卵泡为Driver建立的为反向消减文库。实验过程中,总RNA与反转录成的cDNA质量较好,消减产物得到很好的富集,正向消减文库中获得了79条独立的差异表达基因,BLAST比对结果显示,其中67条具有同源基因,12条无相似基因或者同源性极低的基因；筛选出与卵泡发育相关基因EGFR,与排卵有关的基因ADAMTS-1影响脑神经系统和胚胎发育的编码钙/钙调蛋白依赖性丝氨酸蛋白激酶基因、免疫相关基因lymphocyte antigen86-like和IL17RA,参与细胞调亡的基因death domain-containing tumor necrosis factor receptor superfamily member23,骨成形蛋白2基因。分离了一些编码通道或转运有关蛋白的基因及其同源基因sodium/myo-inositol cotransporter-like、calcium/calmodulin-dependent serine protein kinase、 SLC9A8、potassium inwardly-rectifying channel, subfamily J, member2、 SLC2A1、PCTP等。除上述基因外,还筛选到RNF14、TTC17、PHF21A、PLEKHB2、 ADD3、RGS12、SVEP1、HDAC4、STOM、SMARCE1、SETD4、CEBPG、STRADA、ZC3H14、 RHOBTB1、ALDH1A2、EXOC2和GAPVD等基因。反向消减文库中获得了37条独立的差异表达的基因,比对结果显示,其中29条具有同源基因,8条无相似或相似度极低的基因。反向文库中筛选出的基因主要包括RBMII、微管组分基因α-tubulin lc、 CD58、真核生物翻译起始基因EIF3I以及编码微小染色体维持蛋白的MCM5基因。
The rate of ovulation is determined by, among other things, the availability of small follicles that can be recruited into the follicular hierarchy. A decrease in the rate of lay with, for instance, aging has been attributed to both an increase in the rate of atresia and a decrease in the number of small, growing follicles that provide the pool from which follicles are selected into the final growth phase (the preovulatory hierarchy). Previous studies on follicles of human and mammal ian have shown that the growth and differ entiation of follicles is a complex physiological and biochemical process regulated by many factors, and regulated by not only endocrine hormone of reproductive axis, but also many of the paracrine and autocrine factors secreted by the local ovarian. For research on avian ovarian follicle development, some differentially expressed genes have already been detected in different development stages, but to complex organism is concerned, it is only one small part. Until now, there is still no reasonable hypothesis to illustrate the mechanism of avian hierarchal follicle establishment. Through the screening and comparing of the differentially expressed genes in duck follicles of different development stages, we can furtherly understand the effects of these genes on the process of follicular growth and differentiation, the molecular biology mechanism of the poultry sexual maturity, the start of laying and maintenance of a high yield performance. This study will provide a theoretical foundation for finally revealing the mechanism of avian hierarchal follicle establishment. A better understanding of differentially expressed genes during follicle selection and differentiation should lead to the development of management practices or genetic manipulations (molecular), which could enhance the rate of laying at times during the production cycles when egg production is suboptimal.
In this experiment, we used the follicles of Shaoxing duck as the experimental material, and the DSN mediated normalization subtractive hybridization method to construct two forward-and reverse-substracted cDNA libraries of follicular development-related genes, thus we can find the new genes, and explore the molecular mechanism of avian hierarchal follicle establishment. The results are as follows:
The first forward-and reverse-substracted cDNA libraries between Fl and PF were constructed by SSH technique. In the forward-subs tracted cDNA library, the follicles of F1and PF were used as tester and driver, respectively. Conversely, the reverse library was constructed. Results showed that the qualities total RNA and reverse transcription of cDNA were very good, and the products of SSH were well enr iched. In the forward library, a total of88differentially expressed genes were obtained. After analysis by BLAST, the results showed that60genes have homo logy and the other28genes have no homology from NCBI. Further analysis showed that some important functional genes were isolated in the forward library, including the LHR, ECFR, SLC family, NADH1β, PRKG2and AKAP2, etc. In the reverse-substracted cDNA library, a total of72differentially expressed genes were obtained. After analysis by BLAST, the results showed that51genes have homology and the other21genes have no homology from NCBI. The genes were isolated, including RBMII, p32and U2AF subunit of splicing factor SF2, tensin, EIF3I, α-tubulin, ACTG2, CD58, MCM5, RCC2, Flotillin-2and some other genes encoding enzymes and their homologies, such as FG3ATPase family gene3-like2, AK3L2and TAOK3.
The second forward-and reverse-substracted cDNA libraries between Fl and F6were constructed by SSH technique. In the forward-substracted cDNA library, the follicles of Fl and F6were used as tester and driver, respectively. Conversely, the reverse library was constructed. Resuts showed that the qualities of total RNA and reverse transcription of cDNA were very good, and the products of SSH were well enriched. In the forward library, a total of79differentially expressed genes were obtained. After analysis by BLAST, the results showed that67genes have homology and the other12genes have no homology from NCBI. Further analysis showed that some important functional genes were isolated in the library, including EGFR, ADAMTS-1, MAGUK family, SLC family, lymphocyte antigen86-like, IL17RA, and death domain-containing tumor necrosis factor receptor superfami ly member23. Some genes encoding channel or transporter proteins were isolated, such as sodium/myo-inositol cotransporter-like, calcium/calmodulin-dependent serine protein kinase, SLC9A8, potassium inwardly-rectifying channel, subfamily J, member2. SLC2A1and PCTP, etc. In addition, RNF14, TTC17, PHF21A, PLEKHB2, ADD3, RGS12, SVEP1, HDAC4, STOM, SMARCE1, SETD4, CEBPG, STRADA, ZC3H14, RHOBTB1, ALDH1A2, EXOC2and GAPVD genes were also isolated in this forward library. In the reverse-substracted cDNA library, a total of37differentially expressed genes were obtained. After analysis by BLAST, the results showed that29genes have homology and the other8genes have no homology from NCBI. In this library RBMII, α-tubulin1c, CD58, EIF3I and MCM5genes were isolated.
本实验以产蛋高峰期绍兴鸭母鸭不同阶段的卵泡为实验材料,采用DSN酶介导的均一化消减杂交方法构建鸭卵泡差异表达基因的消减cDNA文库,以期找到与鸭卵泡发育相关的基因,探索卵泡等级建立与分化的基因表达调控的分子机制。本实验构建了两个鸭卵泡差异表达基因的双向消减文库：
以最大等级卵泡和等级前卵泡这两个不同发育阶段的卵泡为实验材料进行消减杂交。其中,以最大等级卵泡为Tester,等级前卵泡为Driver,建立的为正向消减文库；以等级前卵泡为Tester最大等级卵泡为Driver建立的为反向消减文库。实验过程中,总RNA与反转录成的cDNA质量较好,消减产物得到很好的富集。正向消减文库中获得了88条独立的差异表达的基因,BLAST比对结果显示,其中60条具有同源基因,28条相似度极低或无同源性的基因。筛选出与卵泡发育相关基因LHR和EGFR,与细胞增殖和分化有关的基因SLC家族等,影响胚胎发育的NADH1α脱氢酶同源基因NADH1β几个编码激酶或其同源基因如PRKG2、 AKAP2、similar to protein kinase D1。此外,筛选到我们还分离了一些未知功能的基因。反向消减文库中获得了72条独立的差异表达的基因,BLAST比对结果显示,其中51条具有同源基因,21条无相似或者同源性极低的基因。筛选出的基因有RBMII,拼接因子SF2亚单位p32和U2AF35kDa subunit-like, tensin基因,真核生物翻译起始基因EIF3I,微管组分基因α-tubu1in,肠平滑肌肌动蛋白gamma2,含靶蛋白同源基因similar to palladin,免疫反应相关基因CD58,编码微小染色体维持蛋白的基因MCM5, RCC2,参与细胞侵袭和转移的基因Flotillin-2,酶类(包括激酶)及其同源基因AFG3ATPase f ami ly gene3-like2、AK3L2、TAOK3等基因。另外,我们还分离了一些未知功能的新基因。
以最大等级卵泡和最小等级卵泡这两个不同发育阶段的卵泡为实验材料进行消减杂交。其中,以最大等级卵泡为Tester,最小等级卵泡为Driver,建立的为正向消减文库；以最小等级卵泡为Tester,最大等级卵泡为Driver建立的为反向消减文库。实验过程中,总RNA与反转录成的cDNA质量较好,消减产物得到很好的富集,正向消减文库中获得了79条独立的差异表达基因,BLAST比对结果显示,其中67条具有同源基因,12条无相似基因或者同源性极低的基因；筛选出与卵泡发育相关基因EGFR,与排卵有关的基因ADAMTS-1影响脑神经系统和胚胎发育的编码钙/钙调蛋白依赖性丝氨酸蛋白激酶基因、免疫相关基因lymphocyte antigen86-like和IL17RA,参与细胞调亡的基因death domain-containing tumor necrosis factor receptor superfamily member23,骨成形蛋白2基因。分离了一些编码通道或转运有关蛋白的基因及其同源基因sodium/myo-inositol cotransporter-like、calcium/calmodulin-dependent serine protein kinase、 SLC9A8、potassium inwardly-rectifying channel, subfamily J, member2、 SLC2A1、PCTP等。除上述基因外,还筛选到RNF14、TTC17、PHF21A、PLEKHB2、 ADD3、RGS12、SVEP1、HDAC4、STOM、SMARCE1、SETD4、CEBPG、STRADA、ZC3H14、 RHOBTB1、ALDH1A2、EXOC2和GAPVD等基因。反向消减文库中获得了37条独立的差异表达的基因,比对结果显示,其中29条具有同源基因,8条无相似或相似度极低的基因。反向文库中筛选出的基因主要包括RBMII、微管组分基因α-tubulin lc、 CD58、真核生物翻译起始基因EIF3I以及编码微小染色体维持蛋白的MCM5基因。
The rate of ovulation is determined by, among other things, the availability of small follicles that can be recruited into the follicular hierarchy. A decrease in the rate of lay with, for instance, aging has been attributed to both an increase in the rate of atresia and a decrease in the number of small, growing follicles that provide the pool from which follicles are selected into the final growth phase (the preovulatory hierarchy). Previous studies on follicles of human and mammal ian have shown that the growth and differ entiation of follicles is a complex physiological and biochemical process regulated by many factors, and regulated by not only endocrine hormone of reproductive axis, but also many of the paracrine and autocrine factors secreted by the local ovarian. For research on avian ovarian follicle development, some differentially expressed genes have already been detected in different development stages, but to complex organism is concerned, it is only one small part. Until now, there is still no reasonable hypothesis to illustrate the mechanism of avian hierarchal follicle establishment. Through the screening and comparing of the differentially expressed genes in duck follicles of different development stages, we can furtherly understand the effects of these genes on the process of follicular growth and differentiation, the molecular biology mechanism of the poultry sexual maturity, the start of laying and maintenance of a high yield performance. This study will provide a theoretical foundation for finally revealing the mechanism of avian hierarchal follicle establishment. A better understanding of differentially expressed genes during follicle selection and differentiation should lead to the development of management practices or genetic manipulations (molecular), which could enhance the rate of laying at times during the production cycles when egg production is suboptimal.
In this experiment, we used the follicles of Shaoxing duck as the experimental material, and the DSN mediated normalization subtractive hybridization method to construct two forward-and reverse-substracted cDNA libraries of follicular development-related genes, thus we can find the new genes, and explore the molecular mechanism of avian hierarchal follicle establishment. The results are as follows:
The first forward-and reverse-substracted cDNA libraries between Fl and PF were constructed by SSH technique. In the forward-subs tracted cDNA library, the follicles of F1and PF were used as tester and driver, respectively. Conversely, the reverse library was constructed. Results showed that the qualities total RNA and reverse transcription of cDNA were very good, and the products of SSH were well enr iched. In the forward library, a total of88differentially expressed genes were obtained. After analysis by BLAST, the results showed that60genes have homo logy and the other28genes have no homology from NCBI. Further analysis showed that some important functional genes were isolated in the forward library, including the LHR, ECFR, SLC family, NADH1β, PRKG2and AKAP2, etc. In the reverse-substracted cDNA library, a total of72differentially expressed genes were obtained. After analysis by BLAST, the results showed that51genes have homology and the other21genes have no homology from NCBI. The genes were isolated, including RBMII, p32and U2AF subunit of splicing factor SF2, tensin, EIF3I, α-tubulin, ACTG2, CD58, MCM5, RCC2, Flotillin-2and some other genes encoding enzymes and their homologies, such as FG3ATPase family gene3-like2, AK3L2and TAOK3.
The second forward-and reverse-substracted cDNA libraries between Fl and F6were constructed by SSH technique. In the forward-substracted cDNA library, the follicles of Fl and F6were used as tester and driver, respectively. Conversely, the reverse library was constructed. Resuts showed that the qualities of total RNA and reverse transcription of cDNA were very good, and the products of SSH were well enriched. In the forward library, a total of79differentially expressed genes were obtained. After analysis by BLAST, the results showed that67genes have homology and the other12genes have no homology from NCBI. Further analysis showed that some important functional genes were isolated in the library, including EGFR, ADAMTS-1, MAGUK family, SLC family, lymphocyte antigen86-like, IL17RA, and death domain-containing tumor necrosis factor receptor superfami ly member23. Some genes encoding channel or transporter proteins were isolated, such as sodium/myo-inositol cotransporter-like, calcium/calmodulin-dependent serine protein kinase, SLC9A8, potassium inwardly-rectifying channel, subfamily J, member2. SLC2A1and PCTP, etc. In addition, RNF14, TTC17, PHF21A, PLEKHB2, ADD3, RGS12, SVEP1, HDAC4, STOM, SMARCE1, SETD4, CEBPG, STRADA, ZC3H14, RHOBTB1, ALDH1A2, EXOC2and GAPVD genes were also isolated in this forward library. In the reverse-substracted cDNA library, a total of37differentially expressed genes were obtained. After analysis by BLAST, the results showed that29genes have homology and the other8genes have no homology from NCBI. In this library RBMII, α-tubulin1c, CD58, EIF3I and MCM5genes were isolated.
引文
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