在飞蝗两型转变中takeout蛋白功能的研究
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摘要
飞蝗是一种分布广泛的农业害虫,遍布欧、亚、非、澳四大洲。在我国也广泛分布。飞蝗具有两型现象,分为散居型和群居型,且两型在不同环境条件下可以相互转变,散居型向群居型转变并且集聚迁飞是蝗灾爆发的主要原因。
takeout家族是昆虫在长期进化过程中形成的一个特有的、大的蛋白家族,takeout包括大约250个氨基酸,是一类分泌蛋白,主要分布于涉及化学感受和营养的组织,例如果蝇的触角、下唇须、脂肪体及前胃等。其主要功能是参与节律调控,调节昆虫摄食与运动,参与味觉和嗅觉系统的化学感受,参与性别分化,有助于雄性果蝇求偶等。许多学者认为takeout蛋白具有配体结合特征,它会与不同的疏水配体结合从而执行不同的生理功能,但是没有直接证据。
最近有资料表明在群居型飞蝗散居化和散居型飞蝗群居化过程中共有453条共同差异表达的基因,包括多个takeout基因,其中一个takeout基因——LimgTO1被认为通过接收其它个体的化学信号,从而产生应答参与型变过程,但对于其他takeout基因与飞蝗型变的关系尚不清楚。
本研究选取其中一个takeout基因——LimgTO2,以四龄第三天的蝗蝻作为研究对象。首先对其进行信号肽预测,发现其具有信号肽序列;进行进化分析,发现LimgTO2属于takeout家族;进行序列比对分析,发现在N端具有两个保守的Cys残基,推测其可能具有配体结合特征。第二,通过实时荧光定量PCR技术和Western blot方法检测LimgTO2在两型飞蝗的表达,发现散居型飞蝗后腿、翅膀、触角的表达量明显高于群居型,尤其是后腿的差异量最大,而LimgTO2在两型飞蝗的其他组织中均没有表达。第三,同样用实时荧光定量PCR技术和Western blot方法检测检测两型转化过程中的LimgTO2的表达变化,结果显示LimgTO2在散居型飞蝗群居化过程中表达量下降;在群居型飞蝗散居化过程中表达量上升,表明后腿中LimgTO2可能参与两型转变。最后,用实时荧光定量PCR方法、Western blot方法和免疫组织化学方法共同检测,发现LimgTO2位于散居飞蝗的后腿腿节的表皮,推测LimgTO2蛋白可能作为载体蛋白参与后腿表皮感受器感受外界信号,从而进一步影响型变。
国内外许多学者对蝗灾及飞蝗两型转变的机制进行了大量研究,但很少深入到分子生物学层面,本实验从分子生物学角度对飞蝗两型转变的机制进行研究。目前治理蝗灾的方法主要集中于大规模使用化学药物,但这些药物会导致环境污染,生态破坏等问题,而本实验为人类从生物学角度控制蝗灾奠定了基础。
The migratory locust, Locusta migratoria, is a widespread pest that is distributed in Europe, Asia, Africa, and Australia. They also have a broad distribution in China. In L. migratoria, there are two phases, solitarious and gregarious phases, which can transform into the other. Locust outbreaks are mainly caused by the transform of the solitarious into the gregarious phase and the formation of swarm bands.
Takeout family is a big insect-specific protein family formed during long evolution. They are secretary proteins containing about 250 amino acids. Takeout proteins are abundantly distributed in several tissues related to chemical sense and nutrition, for example, antennae, tarsi, fat body and cardia. So far, they are revealed to be involved in circadian rhythm, feeding and locomotor activity, chemical sensation, sex-determination, and male courtship behavior. Many scholars thought that takeout proteins could bind different hydrophobic ligands which could perform different functions, but there were no direct evidence.
Recent research revealed that, during solitarization and gregarization of L. migratoria, the expression of 453 genes differed significantly, including many takeout genes. Among these takeout genes, LimgTO1 is proposed to accept singals from other individuals causing phase change, but the relationship between other takeout genes and phase change remains unclear.
We analyzed the third-day fourth instar nymphs using LimgTO2, which is one of the above takeout gene families. The signal peptide prediction revealed that LimgTO2 possesses signal peptide sequence. Phylogenetic analyses supported that LimgTO2 belongs to the takeout family. The two Cys residues in the N terminus indicated that LimgTO2 could bind ligands. The tissue-specific expression and differential expression profiling of the LimgTO2 gene during phase transition of L. migratoria were analyzed by real-time quantity PCR and Western blot. The gene had a significantly higher expression level in solitarious locusts than gregarious locusts, with the most different expression in the tissue of hind legs. The expression of LimgTO2 was down-regulated during the gregarizaiton process and up-regualted during the reverse process. The above results indicate that the LimgTO2 gene is involved in the regulation of locust phase transition. The results of real-time quantity PCR, Western blot and immunohistochemistry methods reveal that the LimgTO2 protein is distributed in epidermis of locust hind legs. The conclusion is that the sensory hair in hind legs of solitarious locusts can sense external stimuli and transmit the signals via some hormones or signal proteins, and that the LimgTO2 gene acts as a carrying protein involved in the phase transition of L. migratoria.
Although there have been many strudies about the mechanism of the phase transition of the L. migratoria, the present research represents the effort at the molecular level. Moreover, current methods of locust control are mainly focusd on chemical drugs which may cause environmental and ecological pollution. The present study provides solid genetic evidence for the phase transition of L. migratoria, which may help improve the design of biological pest-control measures.
takeout家族是昆虫在长期进化过程中形成的一个特有的、大的蛋白家族,takeout包括大约250个氨基酸,是一类分泌蛋白,主要分布于涉及化学感受和营养的组织,例如果蝇的触角、下唇须、脂肪体及前胃等。其主要功能是参与节律调控,调节昆虫摄食与运动,参与味觉和嗅觉系统的化学感受,参与性别分化,有助于雄性果蝇求偶等。许多学者认为takeout蛋白具有配体结合特征,它会与不同的疏水配体结合从而执行不同的生理功能,但是没有直接证据。
最近有资料表明在群居型飞蝗散居化和散居型飞蝗群居化过程中共有453条共同差异表达的基因,包括多个takeout基因,其中一个takeout基因——LimgTO1被认为通过接收其它个体的化学信号,从而产生应答参与型变过程,但对于其他takeout基因与飞蝗型变的关系尚不清楚。
本研究选取其中一个takeout基因——LimgTO2,以四龄第三天的蝗蝻作为研究对象。首先对其进行信号肽预测,发现其具有信号肽序列;进行进化分析,发现LimgTO2属于takeout家族;进行序列比对分析,发现在N端具有两个保守的Cys残基,推测其可能具有配体结合特征。第二,通过实时荧光定量PCR技术和Western blot方法检测LimgTO2在两型飞蝗的表达,发现散居型飞蝗后腿、翅膀、触角的表达量明显高于群居型,尤其是后腿的差异量最大,而LimgTO2在两型飞蝗的其他组织中均没有表达。第三,同样用实时荧光定量PCR技术和Western blot方法检测检测两型转化过程中的LimgTO2的表达变化,结果显示LimgTO2在散居型飞蝗群居化过程中表达量下降;在群居型飞蝗散居化过程中表达量上升,表明后腿中LimgTO2可能参与两型转变。最后,用实时荧光定量PCR方法、Western blot方法和免疫组织化学方法共同检测,发现LimgTO2位于散居飞蝗的后腿腿节的表皮,推测LimgTO2蛋白可能作为载体蛋白参与后腿表皮感受器感受外界信号,从而进一步影响型变。
国内外许多学者对蝗灾及飞蝗两型转变的机制进行了大量研究,但很少深入到分子生物学层面,本实验从分子生物学角度对飞蝗两型转变的机制进行研究。目前治理蝗灾的方法主要集中于大规模使用化学药物,但这些药物会导致环境污染,生态破坏等问题,而本实验为人类从生物学角度控制蝗灾奠定了基础。
The migratory locust, Locusta migratoria, is a widespread pest that is distributed in Europe, Asia, Africa, and Australia. They also have a broad distribution in China. In L. migratoria, there are two phases, solitarious and gregarious phases, which can transform into the other. Locust outbreaks are mainly caused by the transform of the solitarious into the gregarious phase and the formation of swarm bands.
Takeout family is a big insect-specific protein family formed during long evolution. They are secretary proteins containing about 250 amino acids. Takeout proteins are abundantly distributed in several tissues related to chemical sense and nutrition, for example, antennae, tarsi, fat body and cardia. So far, they are revealed to be involved in circadian rhythm, feeding and locomotor activity, chemical sensation, sex-determination, and male courtship behavior. Many scholars thought that takeout proteins could bind different hydrophobic ligands which could perform different functions, but there were no direct evidence.
Recent research revealed that, during solitarization and gregarization of L. migratoria, the expression of 453 genes differed significantly, including many takeout genes. Among these takeout genes, LimgTO1 is proposed to accept singals from other individuals causing phase change, but the relationship between other takeout genes and phase change remains unclear.
We analyzed the third-day fourth instar nymphs using LimgTO2, which is one of the above takeout gene families. The signal peptide prediction revealed that LimgTO2 possesses signal peptide sequence. Phylogenetic analyses supported that LimgTO2 belongs to the takeout family. The two Cys residues in the N terminus indicated that LimgTO2 could bind ligands. The tissue-specific expression and differential expression profiling of the LimgTO2 gene during phase transition of L. migratoria were analyzed by real-time quantity PCR and Western blot. The gene had a significantly higher expression level in solitarious locusts than gregarious locusts, with the most different expression in the tissue of hind legs. The expression of LimgTO2 was down-regulated during the gregarizaiton process and up-regualted during the reverse process. The above results indicate that the LimgTO2 gene is involved in the regulation of locust phase transition. The results of real-time quantity PCR, Western blot and immunohistochemistry methods reveal that the LimgTO2 protein is distributed in epidermis of locust hind legs. The conclusion is that the sensory hair in hind legs of solitarious locusts can sense external stimuli and transmit the signals via some hormones or signal proteins, and that the LimgTO2 gene acts as a carrying protein involved in the phase transition of L. migratoria.
Although there have been many strudies about the mechanism of the phase transition of the L. migratoria, the present research represents the effort at the molecular level. Moreover, current methods of locust control are mainly focusd on chemical drugs which may cause environmental and ecological pollution. The present study provides solid genetic evidence for the phase transition of L. migratoria, which may help improve the design of biological pest-control measures.
引文
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2.康乐,陈永林.关于蝗虫灭灾减灾对策的探讨.中国减灾, 1992, 2(1): 50-52.
3.朱恩林.中国东亚飞蝗发生与治理.北京,中国农业出版社, 1998, 1-2.
4.赵春晓,崔为正.飞蝗的型变及其调控机理[J]中国植保导刊, 2006, (09).
5.陈永林.蝗虫为什么能爆发成灾.人与生物圈,北京,中国人与生物圈国家委员会, 2005, 03: 20-23.
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