蜜蜂保幼激素终端合成相关酶基因分子克隆、鉴定及其在级型发育过程中的表达研究
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摘要
蜜蜂Apis mellifera作为高度真社会性昆虫,已成为社会生物学研究的模式生物。社会性昆虫的生殖劳动分工具有重要的进化意义,而级型分化是形成生殖劳动分工的基础。已有的研究成果为人们描绘出一个相对完善的级型分化调控网络。然而,仍有许多问题值得进一步探索,例如：影响级型分化各分表型——个体大小、发育时长、卵巢大小——的具体分子机制；保幼激素等重要内分泌因子的产生和作用的具体分子途径等。本研究借助新一代高通量测序技术,检测了蜜蜂蜂王和工蜂幼虫级型发育过程基因表达谱。同时,克隆和鉴定了蜜蜂保幼激素(juvenile hormone, JH)生物合成终端步骤相关3种酶基因——保幼激素酸甲基转移酶基因AmJHAMT、细胞色素P45015A1基因AmCYP15A1和法尼酸甲基转移酶基因AmFAMeT,并结合蜜蜂级型发育过程分析了这3个基因可能发挥的功能。主要研究结果如下：
1.基于新一代测序技术的蜜蜂级型分化数字化基因表达谱
本研究采用基于Illumina/Solexa高通量测序技术的数字化基因表达谱方法(digital gene expression profile, DGE),检测了西方蜜蜂蜂王和工蜂幼虫级型发育表达谱。分别从蜂王和工蜂两个DGE测序文库中获得了约760万和890万的可靠标签,相应的独立标签数目分别约为8.7万和10.0万。差异基因表达分析揭示,在蜂王幼虫中分别有1278个显著上调表达基因以及1451个显著下调表达基因,在最显著上调的50个基因中,多数是代谢酶类基因,同时还包括2个参与保幼激素合成的基因——-JHAMT和CYP15A1.相对地,工蜂幼虫最显著上调的基因仅有少数是代谢酶类基因。有62,417个无法比对到参考标签数据库的独立标签唯一比对到蜜蜂基因组序列中,运用Genscan对它们进行基因预测,获得了9,258个候选基因,其中有3,566个候选基因长度超过300nt。GO分析揭示这些候选基因可能参与胚后发育、生殖发育和有性生殖等生理过程。
2.蜜蜂JHAMT酶基因分子克隆、鉴定及其在级型分化中的功能研究
保幼激素酸甲基转移酶(juvenile hormone acid methyltransferase, JHAMT)是一种参与昆虫体内保幼激素终端合成步骤的酶,它将来源于S-腺苷-L-甲硫氨酸(S-adenosyl-L-methionine, SAM)的甲基转移到法尼酸(farnesoic acid, FA)(或保幼激素酸,juvenile hormone acid,JHA)的羧基上。运用RACE技术克隆了西方蜜蜂.JHAMT基因——AmJHAMT的全长cDNA,其长度为1253bp,编码一个278aa的蛋白质,该蛋白与已知的JHAMT蛋白有32%-36%的序列一致性.AmJHAMT包含在SAM依赖性甲基转移酶超家族中保守的SAM结合基序——“基序I”。它的二级结构同样包含SAM甲基转移酶超家族成员典型的核心折叠。同时,那些与SAM及酶促底物——FA或JHA——结合的活性位点在AmJHAMT序列中仍然保守。在大肠杆菌中表达了AmJHAMT蛋白,并用重组蛋白制备了相应的多克隆抗体；运用免疫印迹和质谱技术进一步验证了预测的AmJHAMT蛋白氨基酸序列的正确性。荧光定量PCR和免疫印迹分析显示,在级型发育过程中,蜂王比工蜂表达更多的AmJHAMT mRNA和蛋白质；同时,蜂王与工蜂的AmJHAMT mRNA和蛋白质时序表达谱与JH滴度变化保持相似的波动模式。
3.蜜蜂CYP15A1酶基因分子克隆、鉴定及其在级型分化过程中的表达
细胞色素P450超家族(cytochrome P450superfamily, CYP)的CYP15家族成员(如CYP15A1)催化昆虫JH终端合成步骤的环氧化反应(oxidation),使甲基法尼酯(methyl farnesoate, MF)转变为JH。本研究从西方蜜蜂幼虫中克隆了包含完整ORF的CYP15A1基因——AmCYP15A1,其ORF长为1515bp,编码含504aa的蛋白。多重比对分析发现,AmCYP15A1与多数昆虫CYP15序列一致性低于50%,然而AmCYP15A1以及其他昆虫CYP15蛋白都含有构成CYP蛋白一般结构折叠相应的基序,以及N端膜锚及铰链等CYPs特征结构。系统发生分析显示,AmCYP15A1先后与来自膜翅目、双翅目、鞘翅目、直翅目、蜚蠊目、半翅目以及虱目昆虫的CYP15s聚集成为一个主进化枝,而鳞翅目昆虫CYP15s独立构成另一个主进化枝。在级型发育过程的绝大多数时期,蜂王中的AmCYP15A1mRNA表达丰度显著高于工蜂,尤其是在级型发育的关键窗口期(也称JH敏感期)。另一方面,从表达趋势上来看,无论是在蜂王,还是在工蜂中,AmCYP15A1基因表达丰度均在幼虫L5F时期达到峰值,随后迅速下降,直至Pw期呈现很低的表达水平,这与末龄期-蛹期过程JH滴度变化保持一致。
4.蜜蜂FAMeT酶基因分子克隆、鉴定及其在级型发育过程中的表达
法尼酸甲基转移酶(Farnesoic acid O-methyltransferase, FAMeT)被认为参与JH终端合成过程——FAMeT以SAM为辅助因子催化FA转变为MF。本研究从西方蜜蜂幼虫中成功克隆得到FAMeT基因——AmFAMeT,其cDNA ORF长为891bp,编码含296aa蛋白。多重比对分析表明, AmFAMeT与来自昆虫的FAMeT序列具有更高的一致性(54%-85%),而与甲壳动物的序列一致性较低(<50%)。比较AmFAMeT基因在蜂王和工蜂幼虫相同发育阶段的表达丰度可知,在多数情况下,二者AmFAMeT表达水平无显著差异,同时,基因表达趋势互不一致,且与JH滴度变化也无明显一致性。
综上所述,蜜蜂级型分化是一个伴随着剧烈基因差异表达的过程。AmJHAMT与AmCYP15A1分别是蜜蜂中真实存在的JHAMT酶基因及CYP15酶基因,它们通过调控JH终端生物合成而参与级型分化调节。AmFAMeT与蜜蜂级型分化无关联,可能已失去了催化JH合成的功能。
As a highly eusocial insect, the honey bee, Apis mellifera, has become a model organism in sociobiology. The reproductive division of labor in social insects, based on caste differentiation, is of great significance in evolution. People have been able to build a relatively complete regulatory network of honey bee caste differentiation with the exiting results. However, there are still many issues worthy of further exploration, such as the molecular mechanisms determining the expression of sub-phenotypes like body size, developmental duration, and ovary size, as well as the molecular pathways involved in the biosynthesis and actions of the important endocrine factors such as Juvenile hormone (JH). In this study, the next-generation sequencing (NGS) technology was employed to detect the gene expression profiles of queen and worker caste development. Three genes AmJHAMT, AmCYP15A1and AmFAMeT involved in the final steps of JH biosynthesis in honey bees were cloned and characterized, and their putative functions for honey bee caste development were investigated. The main findings were as follows:
1. Next-generation sequencing-based digital gene expression profiles during honeybee caste differentiation
In this study, the digital gene expression profiling (DGE) was used to detect gene expression profiles responding to the queen and worker caste development. About7.6million and8.9million clean tags, with87,000and100,000distinct tags were obtained from the queen and worker DGE sequencing library, respectively.1,278and1,451genes were found to be significantly up-regulated and down-regulated in queen larvae compared with worker larvae. The majority of the top50significantly up-regulated genes in queen larvae were metabolic enzymes, and two juvenile hormone synthesis-related genes, JHAMT and CYP15A1were also included in this set. In contrast, only a few metabolic enzymes were detected in the50most significantly up-regulated gene list of worker larvae. There were62,417entities uniquely mapped to the honey bee genome, which failed to map to the reference tag database, and9,258candidate genes were obtained by gene prediction from these entities using Genscan program, of which3,566candidate genes were longer than300nt. GO analysis revealed that these candidate genes might be involved in the embryonic development, reproductive development and sexual reproduction.
2. Molecular cloning and characterization of JHAMT in A. mellifera, and its functions during caste differentiation
Juvenile hormone acid methyltransferase (JHAMT) is an enzyme involved in one of the final steps of juvenile hormone biosynthesis in insects, which transfers a methyl group from S-adenosyl-L-methionine (SAM) to the carboxyl group of either farnesoic acid (FA) or JH acid (JHA). RACE amplification method was used to clone JHAMT cDNA from the honey bee, Apis mellifera(AmJHAMT). The full length cDNA of AmJHAMT was1253bp long and encoded a278-aa protein that shared32%-36%identity with known JHAMTs. A SAM-binding motif, conserved in the SAM-dependent methyltransferase (SAM-MT) superfamily, was present in AmJHAMT. Its secondary structure also contained a typical SAM-MT fold. Most of the active sites bound with SAM and substrates (JHA or FA) were conserved in AmJHAMT as in other JHAMT orthologs. Purified recombinant AmJHAMT protein expressed in E. coli was used to produce polyclonal antibodies, and the identity of AmJHAMT was verified latter by immunoblotting and mass spectrometry. Quantitative RT-PCR and immunoblotting analyses revealed that queen larvae contained significantly higher levels of AmJHAMT mRNA and protein than worker larvae during the periods of caste development. The temporal profiles of both AmJHAMT mRNA and protein in queens and workers showed a similar pattern as the JH biosynthesis.
3. Molecular cloning and characterization of CYP15A1in A. mellifera, and its expression during caste differentiation
The CYP15family members like CYP15A1of Cytochrome P450superfamily (CYP) catalyze the epoxidation reaction of the final steps of JH biosynthesis in insects, which transfers methyl farnesoate (MF) to JH. In this study, a putative CYP15A1gene, named AmCYP15A1, was cloned from the queen larvae of A. mellifera. It contained an1515bp ORF, which encoded an504aa protein. Multiple alignment analysis showed a relatively low identity between AmCYP15A1with most other insect CYP15s (<50%). However, AmCYP15A1and the other insect CYP15s all contained the motifs constituting the general CYP structure fold, and some other characteristic structures of CYPs, like N-terminal membrane anchor and hinges and so on. Phylogenetic analysis displayed that AmCYP15A1was successively clustered with the CYP15orthologs of insects from Hymenoptera, Diptera, Coleoptera, Orthoptera, Blattodea, and Hemiptera, which finally formed a major clade. However, the other major clade was constituted only by Lepidopteran CYP15s. The gene expression level of AmCYP15A1was determined by qPCR, and the result showed that the AmCYP15A1mRNA expression levels in queens were significantly higher than those in workers during most of developmental stages, especially for the physiologically critical temporal window of caste differentiation (also known as JH-sensitive period). On the other hand, the AmCYP15A1mRNA levels of both queens and workers peaked in the feeding stage of the5th instar, and then decreased rapidly to a low level until the white-eyed pupal stage, which was consist with JH biosynthesis activity.
4. Molecular cloning and characterization of FAMeT in A. mellifera, and its expression during caste development
Farnesoic acid methyl transferases (FAMeT) are thought to be involved in the terminal steps of JH biosynthesis, which catalyzes the methylation of FA to MF using the cofactor SAM. In this study, a putative FAMeT gene, namely AmFAMeT, was cloned from the queen larvae of western honey bees. It contained an891bp ORF which encoded an504aa protein. Multiple alignment analysis indicated that AmFAMeT had a higher identity with insects FAMeTs (54%-85%) when compared with crustacean ones (<50%). Comparison of AmFAMeT expression abundance between queen and workers showed that there were no significant differences for the most of caste developmental stages. Meanwhile, the trends of gene expression differed between the two castes, and were both not consist with the JH biosynthesis.
In a word, our results suggest that:1) Honey bee caste differentiation is a process with significantly differential gene expression;2) AmJHAMT and AmCYP15A1are real JHAMT and CYP15enzyme genes in A. mellifera, which involve in the regulation of caste differentiation by control the terminal biosynthesis of JH;3) AmFAMeT is not related to the caste development, and may lose the function of catalyzing final reactions of JH biosynthesis.
1.基于新一代测序技术的蜜蜂级型分化数字化基因表达谱
本研究采用基于Illumina/Solexa高通量测序技术的数字化基因表达谱方法(digital gene expression profile, DGE),检测了西方蜜蜂蜂王和工蜂幼虫级型发育表达谱。分别从蜂王和工蜂两个DGE测序文库中获得了约760万和890万的可靠标签,相应的独立标签数目分别约为8.7万和10.0万。差异基因表达分析揭示,在蜂王幼虫中分别有1278个显著上调表达基因以及1451个显著下调表达基因,在最显著上调的50个基因中,多数是代谢酶类基因,同时还包括2个参与保幼激素合成的基因——-JHAMT和CYP15A1.相对地,工蜂幼虫最显著上调的基因仅有少数是代谢酶类基因。有62,417个无法比对到参考标签数据库的独立标签唯一比对到蜜蜂基因组序列中,运用Genscan对它们进行基因预测,获得了9,258个候选基因,其中有3,566个候选基因长度超过300nt。GO分析揭示这些候选基因可能参与胚后发育、生殖发育和有性生殖等生理过程。
2.蜜蜂JHAMT酶基因分子克隆、鉴定及其在级型分化中的功能研究
保幼激素酸甲基转移酶(juvenile hormone acid methyltransferase, JHAMT)是一种参与昆虫体内保幼激素终端合成步骤的酶,它将来源于S-腺苷-L-甲硫氨酸(S-adenosyl-L-methionine, SAM)的甲基转移到法尼酸(farnesoic acid, FA)(或保幼激素酸,juvenile hormone acid,JHA)的羧基上。运用RACE技术克隆了西方蜜蜂.JHAMT基因——AmJHAMT的全长cDNA,其长度为1253bp,编码一个278aa的蛋白质,该蛋白与已知的JHAMT蛋白有32%-36%的序列一致性.AmJHAMT包含在SAM依赖性甲基转移酶超家族中保守的SAM结合基序——“基序I”。它的二级结构同样包含SAM甲基转移酶超家族成员典型的核心折叠。同时,那些与SAM及酶促底物——FA或JHA——结合的活性位点在AmJHAMT序列中仍然保守。在大肠杆菌中表达了AmJHAMT蛋白,并用重组蛋白制备了相应的多克隆抗体；运用免疫印迹和质谱技术进一步验证了预测的AmJHAMT蛋白氨基酸序列的正确性。荧光定量PCR和免疫印迹分析显示,在级型发育过程中,蜂王比工蜂表达更多的AmJHAMT mRNA和蛋白质；同时,蜂王与工蜂的AmJHAMT mRNA和蛋白质时序表达谱与JH滴度变化保持相似的波动模式。
3.蜜蜂CYP15A1酶基因分子克隆、鉴定及其在级型分化过程中的表达
细胞色素P450超家族(cytochrome P450superfamily, CYP)的CYP15家族成员(如CYP15A1)催化昆虫JH终端合成步骤的环氧化反应(oxidation),使甲基法尼酯(methyl farnesoate, MF)转变为JH。本研究从西方蜜蜂幼虫中克隆了包含完整ORF的CYP15A1基因——AmCYP15A1,其ORF长为1515bp,编码含504aa的蛋白。多重比对分析发现,AmCYP15A1与多数昆虫CYP15序列一致性低于50%,然而AmCYP15A1以及其他昆虫CYP15蛋白都含有构成CYP蛋白一般结构折叠相应的基序,以及N端膜锚及铰链等CYPs特征结构。系统发生分析显示,AmCYP15A1先后与来自膜翅目、双翅目、鞘翅目、直翅目、蜚蠊目、半翅目以及虱目昆虫的CYP15s聚集成为一个主进化枝,而鳞翅目昆虫CYP15s独立构成另一个主进化枝。在级型发育过程的绝大多数时期,蜂王中的AmCYP15A1mRNA表达丰度显著高于工蜂,尤其是在级型发育的关键窗口期(也称JH敏感期)。另一方面,从表达趋势上来看,无论是在蜂王,还是在工蜂中,AmCYP15A1基因表达丰度均在幼虫L5F时期达到峰值,随后迅速下降,直至Pw期呈现很低的表达水平,这与末龄期-蛹期过程JH滴度变化保持一致。
4.蜜蜂FAMeT酶基因分子克隆、鉴定及其在级型发育过程中的表达
法尼酸甲基转移酶(Farnesoic acid O-methyltransferase, FAMeT)被认为参与JH终端合成过程——FAMeT以SAM为辅助因子催化FA转变为MF。本研究从西方蜜蜂幼虫中成功克隆得到FAMeT基因——AmFAMeT,其cDNA ORF长为891bp,编码含296aa蛋白。多重比对分析表明, AmFAMeT与来自昆虫的FAMeT序列具有更高的一致性(54%-85%),而与甲壳动物的序列一致性较低(<50%)。比较AmFAMeT基因在蜂王和工蜂幼虫相同发育阶段的表达丰度可知,在多数情况下,二者AmFAMeT表达水平无显著差异,同时,基因表达趋势互不一致,且与JH滴度变化也无明显一致性。
综上所述,蜜蜂级型分化是一个伴随着剧烈基因差异表达的过程。AmJHAMT与AmCYP15A1分别是蜜蜂中真实存在的JHAMT酶基因及CYP15酶基因,它们通过调控JH终端生物合成而参与级型分化调节。AmFAMeT与蜜蜂级型分化无关联,可能已失去了催化JH合成的功能。
As a highly eusocial insect, the honey bee, Apis mellifera, has become a model organism in sociobiology. The reproductive division of labor in social insects, based on caste differentiation, is of great significance in evolution. People have been able to build a relatively complete regulatory network of honey bee caste differentiation with the exiting results. However, there are still many issues worthy of further exploration, such as the molecular mechanisms determining the expression of sub-phenotypes like body size, developmental duration, and ovary size, as well as the molecular pathways involved in the biosynthesis and actions of the important endocrine factors such as Juvenile hormone (JH). In this study, the next-generation sequencing (NGS) technology was employed to detect the gene expression profiles of queen and worker caste development. Three genes AmJHAMT, AmCYP15A1and AmFAMeT involved in the final steps of JH biosynthesis in honey bees were cloned and characterized, and their putative functions for honey bee caste development were investigated. The main findings were as follows:
1. Next-generation sequencing-based digital gene expression profiles during honeybee caste differentiation
In this study, the digital gene expression profiling (DGE) was used to detect gene expression profiles responding to the queen and worker caste development. About7.6million and8.9million clean tags, with87,000and100,000distinct tags were obtained from the queen and worker DGE sequencing library, respectively.1,278and1,451genes were found to be significantly up-regulated and down-regulated in queen larvae compared with worker larvae. The majority of the top50significantly up-regulated genes in queen larvae were metabolic enzymes, and two juvenile hormone synthesis-related genes, JHAMT and CYP15A1were also included in this set. In contrast, only a few metabolic enzymes were detected in the50most significantly up-regulated gene list of worker larvae. There were62,417entities uniquely mapped to the honey bee genome, which failed to map to the reference tag database, and9,258candidate genes were obtained by gene prediction from these entities using Genscan program, of which3,566candidate genes were longer than300nt. GO analysis revealed that these candidate genes might be involved in the embryonic development, reproductive development and sexual reproduction.
2. Molecular cloning and characterization of JHAMT in A. mellifera, and its functions during caste differentiation
Juvenile hormone acid methyltransferase (JHAMT) is an enzyme involved in one of the final steps of juvenile hormone biosynthesis in insects, which transfers a methyl group from S-adenosyl-L-methionine (SAM) to the carboxyl group of either farnesoic acid (FA) or JH acid (JHA). RACE amplification method was used to clone JHAMT cDNA from the honey bee, Apis mellifera(AmJHAMT). The full length cDNA of AmJHAMT was1253bp long and encoded a278-aa protein that shared32%-36%identity with known JHAMTs. A SAM-binding motif, conserved in the SAM-dependent methyltransferase (SAM-MT) superfamily, was present in AmJHAMT. Its secondary structure also contained a typical SAM-MT fold. Most of the active sites bound with SAM and substrates (JHA or FA) were conserved in AmJHAMT as in other JHAMT orthologs. Purified recombinant AmJHAMT protein expressed in E. coli was used to produce polyclonal antibodies, and the identity of AmJHAMT was verified latter by immunoblotting and mass spectrometry. Quantitative RT-PCR and immunoblotting analyses revealed that queen larvae contained significantly higher levels of AmJHAMT mRNA and protein than worker larvae during the periods of caste development. The temporal profiles of both AmJHAMT mRNA and protein in queens and workers showed a similar pattern as the JH biosynthesis.
3. Molecular cloning and characterization of CYP15A1in A. mellifera, and its expression during caste differentiation
The CYP15family members like CYP15A1of Cytochrome P450superfamily (CYP) catalyze the epoxidation reaction of the final steps of JH biosynthesis in insects, which transfers methyl farnesoate (MF) to JH. In this study, a putative CYP15A1gene, named AmCYP15A1, was cloned from the queen larvae of A. mellifera. It contained an1515bp ORF, which encoded an504aa protein. Multiple alignment analysis showed a relatively low identity between AmCYP15A1with most other insect CYP15s (<50%). However, AmCYP15A1and the other insect CYP15s all contained the motifs constituting the general CYP structure fold, and some other characteristic structures of CYPs, like N-terminal membrane anchor and hinges and so on. Phylogenetic analysis displayed that AmCYP15A1was successively clustered with the CYP15orthologs of insects from Hymenoptera, Diptera, Coleoptera, Orthoptera, Blattodea, and Hemiptera, which finally formed a major clade. However, the other major clade was constituted only by Lepidopteran CYP15s. The gene expression level of AmCYP15A1was determined by qPCR, and the result showed that the AmCYP15A1mRNA expression levels in queens were significantly higher than those in workers during most of developmental stages, especially for the physiologically critical temporal window of caste differentiation (also known as JH-sensitive period). On the other hand, the AmCYP15A1mRNA levels of both queens and workers peaked in the feeding stage of the5th instar, and then decreased rapidly to a low level until the white-eyed pupal stage, which was consist with JH biosynthesis activity.
4. Molecular cloning and characterization of FAMeT in A. mellifera, and its expression during caste development
Farnesoic acid methyl transferases (FAMeT) are thought to be involved in the terminal steps of JH biosynthesis, which catalyzes the methylation of FA to MF using the cofactor SAM. In this study, a putative FAMeT gene, namely AmFAMeT, was cloned from the queen larvae of western honey bees. It contained an891bp ORF which encoded an504aa protein. Multiple alignment analysis indicated that AmFAMeT had a higher identity with insects FAMeTs (54%-85%) when compared with crustacean ones (<50%). Comparison of AmFAMeT expression abundance between queen and workers showed that there were no significant differences for the most of caste developmental stages. Meanwhile, the trends of gene expression differed between the two castes, and were both not consist with the JH biosynthesis.
In a word, our results suggest that:1) Honey bee caste differentiation is a process with significantly differential gene expression;2) AmJHAMT and AmCYP15A1are real JHAMT and CYP15enzyme genes in A. mellifera, which involve in the regulation of caste differentiation by control the terminal biosynthesis of JH;3) AmFAMeT is not related to the caste development, and may lose the function of catalyzing final reactions of JH biosynthesis.
引文
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