昆虫鞘脂质代谢途径相关酶的生化特征和功能
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摘要
中性神经酰胺酶和中性鞘磷脂酶是生物体体内调节鞘脂质代谢的重要酶类。中性鞘磷脂酶水解神经鞘磷脂生成神经神经酰胺,中性神经酰胺酶水解神经酰胺生成神经鞘氨醇。本研究以赤拟谷盗和灰飞虱为研究对象,对赤拟谷盗和灰飞虱的中性神经酰胺酶、灰飞虱的中性鞘磷脂酶生化特征进行了研究,并且探索了它们在赤拟谷盗和灰飞虱体内可能的功能。以下是本研究的主要结果：
1.从赤拟谷盗基因组中的中性神经酰胺酶的三个同源基因中筛选得到了赤拟谷盗的中性神经酰胺酶Tncer。Tncer蛋白序列和果蝇、水稻、人类、铜绿假单胞杆菌的中性神经酰胺酶蛋白序列的相似性分别为49.1%、38.1%、36.8%和33.4%；Tncer表现了其独特的生化特性：虽然中性、碱性、酸性神经酰胺酶的划分是根据酶发挥最大活性的pH值不同来划分,但是越来越多的研究表明中性神经酰胺酶在最适pH不仅仅是在中性的pH范围内,Tncer的最适pH为pH5,但是它在pH5～12范围的活性差别不大,表明Tncer可以在酸性、中性和碱性的pH环境中都发挥很高的活性；Tncer对底物的选择没有专一性,它可以水解从短链到超长链的神经酰胺,但是对短链神经酰胺如C6-Ceramide的水解能力较低；Tncer发挥体外活性的最适温度为37℃；它不依赖二价阳离子来发挥活性,但是活性可以被Fe2+抑制；Tncer在精巢和卵巢中的表达量最大,成虫羽化后在卵巢的成熟过程中Tncer的表达水平表现出升高的趋势,说明Tncer在卵子的形成过程中很可能发挥着重要的作用；Tncer在粉纹夜蛾细胞中主要分布在细胞膜上,这和哺乳动物中性神经酰胺酶的亚细胞定位是相同的。
2.从灰飞虱转录组数据库中得到中性神经酰胺酶的序列。灰飞虱中性神经酰胺酶LsnCer和赤拟谷盗、果蝇、人类、铜绿假单胞杆菌、水稻的中性神经酰胺酶的同源性分别为50.62%、47.46%、37.11%、36.14%、36.84%；LsnCer的最适pH为8,在强酸和强碱性的环境中几乎没有活性；LsnCer对底物的选择也没有专一性,它偏好中长链的神经酰胺；对短链和超长链的神经酰胺水解能力较弱；LsnCer能够更有效地水解含不饱和碳链鞘氨醇基团的神经酰胺。LsnCer的活性可以被和Zn2+显著抑制；其体外活性的最适温度也是37℃；LsnCer在灰飞虱的精巢和卵巢中的表达量最高；它在粉纹夜蛾细胞中的也主要位于细胞膜上
3.携带水稻条纹叶枯病病毒的灰飞虱体内有较高的中性神经酰胺酶表达量,且在pH8的条件下神经酰胺酶活性较高。分别用吡虫啉、氟虫腈和毒死蜱滴定灰飞虱四龄若虫后发现LsnCer的表达水平在4小时后显著上升,并且酶活水平在4小时后也显著上升。
4.在灰飞虱转录组数据库中拼接得到中性鞘磷脂酶1的同源序列LsnSMase,将LsnSMase和其它已知的中性鞘磷脂酶1做序列联配发现,一些氨基酸在这些中性鞘磷脂酶中是极为保守的；LsnSMase蛋白的C端323-342AA和355-374AA为两个跨膜区域；LsnSMase的活性依赖Mg2+,DTT能够提高LsnSMase的活性,EGTA能够显著提高LsnSMase的活性,但是同为离子螯合剂的EDTA能够显著抑制LsnSMase的活性,LsnSMase的活性能够完全被Zn2+抑制。同时加入Mg2+, DTT或Mg2+,EGTA或Mg2+,EDTA都能够极其显著的提高LsnSMase的活性；LsnSMase在灰飞虱雌虫的卵巢中的表达量最高。
本研究首次对农业害虫的鞘脂质代谢途径进行了探索,明确了中性神经酰胺酶和中性鞘磷脂酶的详细生化特征,分析了它们在生物体内可能具有的功能。
Neutral ceramidase and neutral sphingomyelinase are key enzymes in sphingolipid metabolism. Neutral sphingomyelinase hydrolyzes sphingomyelin to generate ceramide. Neutral ceramidase hydrolyzes ceramide, forming sphingosine. We focused on the sphingolipid metabolism of model insect Tribolium castaneum and rice pest the small brown planthopper Laodelphax striatellus. In these researches, we studied these two enzymes of Tribolium castaneum and Laodelphax striatellus. The main results we got were:
1. We identified the neutral ceramidase of Tribolium castaneum (Tncer) from three candidate homologue genes. The amino acid sequence of Tncer has49.1%,38.1%,36.8%,33.4%similarities to the neutral ceramidase of Drosophila melanogaster, rice, human and Pseudomonas aeruginosa. Tncer showed unique biochemical properties. The optimum pH for Tncer activity is about pH5. From pH5to pH12, the ceramidase activity did not change too much. Tncer could hydrolyze various ceramidase from C6-ceramide to C24:1-ceramide. However the hydrolyzed ability to C6-ceramide is lower. The best temperature for its in vitro activity is about37℃. Tncer is a cation independent enzyme, Fe2+can inhibit its activity. The mRNA level of Tncer is highest in spermary and ovary of Tribolium castaneum. And the mRNA level of Tncer is increased with the maturation of ovary. These suggest that Tncer may paly roles in the oogenesis. Tncer is mainly located in the plasma membrane in High Five cells.
2. With the sequence assembly, we got the neutral ceramidase homologue (LsnCer) from the transcriptome database of Laodelphax striatellus. The sequence alignment showed that the sequence similarities between LsnCer and neutral ceramidase of Tribolium castaneum, Drosophila melanogaster, human, Pseudomonas aeruginosa and rice is50.62%,47.76%,37.11%,36.14%and36.84%respectively. The optimum pH for LsnCer is pH8. Little activity was detected in the strong acid and alkaline pH environment. It prefers medium-chain to long-chain ceramide as substrates. Short chain-ceramides such as C2-ceramide and super long chain-ceramide are resistant to LsnCer. The activity of LsnCer could be inhibited significantly by Zn2+The best temperature for its in vitro activity is also37℃. The mRNA level of LsnCer is highest in spermary and ovary of Laodelphax striatellus. It's a transmembrane protein and located in the plasma membrane in the High Five cells.
3. Rice stripe virus (RSV) can stimulate the expression of LsnCer. The ceramidase activity of LsnCer around pH8is also higher in RSV-infected Laodelphax striatellus. mRNA level of LsnCer as well as the neutral ceramidase activity were stimulated significantly after treated with insecticide4hours later.
4. We cloned neutral ceramidase1homologue (LsnSMase) from Laodelphax striatellus.323-342AA and355-374AA are two transmembrane domains. With the bioinformatic analysis and reported results, we presumed LsnSMase is the main sphingomyelinase in Laodelphax striatellus. The activity of LsnSMase is dependent on Mg2+, DTT can also increased the activity of LsnSMase. EDTA inhibited LsnSMase activity significantly. However, EGTA is better stimulator of LsnSMase than Mg2+. The co-operation of Mg2+and EGTA or Mg2+and EDTA or Mg2+and DTT could increase the LsnSMase activity sharply. mRNA level of LsnSMase is highest in ovary of Laodelphax striatellus.
In conclusion, we first identified biochemical properties of neutral ceramidase and neutral sphingomyelinase of agriculture pest. The function of these enzymes are also uncovered in our research.
1.从赤拟谷盗基因组中的中性神经酰胺酶的三个同源基因中筛选得到了赤拟谷盗的中性神经酰胺酶Tncer。Tncer蛋白序列和果蝇、水稻、人类、铜绿假单胞杆菌的中性神经酰胺酶蛋白序列的相似性分别为49.1%、38.1%、36.8%和33.4%；Tncer表现了其独特的生化特性：虽然中性、碱性、酸性神经酰胺酶的划分是根据酶发挥最大活性的pH值不同来划分,但是越来越多的研究表明中性神经酰胺酶在最适pH不仅仅是在中性的pH范围内,Tncer的最适pH为pH5,但是它在pH5～12范围的活性差别不大,表明Tncer可以在酸性、中性和碱性的pH环境中都发挥很高的活性；Tncer对底物的选择没有专一性,它可以水解从短链到超长链的神经酰胺,但是对短链神经酰胺如C6-Ceramide的水解能力较低；Tncer发挥体外活性的最适温度为37℃；它不依赖二价阳离子来发挥活性,但是活性可以被Fe2+抑制；Tncer在精巢和卵巢中的表达量最大,成虫羽化后在卵巢的成熟过程中Tncer的表达水平表现出升高的趋势,说明Tncer在卵子的形成过程中很可能发挥着重要的作用；Tncer在粉纹夜蛾细胞中主要分布在细胞膜上,这和哺乳动物中性神经酰胺酶的亚细胞定位是相同的。
2.从灰飞虱转录组数据库中得到中性神经酰胺酶的序列。灰飞虱中性神经酰胺酶LsnCer和赤拟谷盗、果蝇、人类、铜绿假单胞杆菌、水稻的中性神经酰胺酶的同源性分别为50.62%、47.46%、37.11%、36.14%、36.84%；LsnCer的最适pH为8,在强酸和强碱性的环境中几乎没有活性；LsnCer对底物的选择也没有专一性,它偏好中长链的神经酰胺；对短链和超长链的神经酰胺水解能力较弱；LsnCer能够更有效地水解含不饱和碳链鞘氨醇基团的神经酰胺。LsnCer的活性可以被和Zn2+显著抑制；其体外活性的最适温度也是37℃；LsnCer在灰飞虱的精巢和卵巢中的表达量最高；它在粉纹夜蛾细胞中的也主要位于细胞膜上
3.携带水稻条纹叶枯病病毒的灰飞虱体内有较高的中性神经酰胺酶表达量,且在pH8的条件下神经酰胺酶活性较高。分别用吡虫啉、氟虫腈和毒死蜱滴定灰飞虱四龄若虫后发现LsnCer的表达水平在4小时后显著上升,并且酶活水平在4小时后也显著上升。
4.在灰飞虱转录组数据库中拼接得到中性鞘磷脂酶1的同源序列LsnSMase,将LsnSMase和其它已知的中性鞘磷脂酶1做序列联配发现,一些氨基酸在这些中性鞘磷脂酶中是极为保守的；LsnSMase蛋白的C端323-342AA和355-374AA为两个跨膜区域；LsnSMase的活性依赖Mg2+,DTT能够提高LsnSMase的活性,EGTA能够显著提高LsnSMase的活性,但是同为离子螯合剂的EDTA能够显著抑制LsnSMase的活性,LsnSMase的活性能够完全被Zn2+抑制。同时加入Mg2+, DTT或Mg2+,EGTA或Mg2+,EDTA都能够极其显著的提高LsnSMase的活性；LsnSMase在灰飞虱雌虫的卵巢中的表达量最高。
本研究首次对农业害虫的鞘脂质代谢途径进行了探索,明确了中性神经酰胺酶和中性鞘磷脂酶的详细生化特征,分析了它们在生物体内可能具有的功能。
Neutral ceramidase and neutral sphingomyelinase are key enzymes in sphingolipid metabolism. Neutral sphingomyelinase hydrolyzes sphingomyelin to generate ceramide. Neutral ceramidase hydrolyzes ceramide, forming sphingosine. We focused on the sphingolipid metabolism of model insect Tribolium castaneum and rice pest the small brown planthopper Laodelphax striatellus. In these researches, we studied these two enzymes of Tribolium castaneum and Laodelphax striatellus. The main results we got were:
1. We identified the neutral ceramidase of Tribolium castaneum (Tncer) from three candidate homologue genes. The amino acid sequence of Tncer has49.1%,38.1%,36.8%,33.4%similarities to the neutral ceramidase of Drosophila melanogaster, rice, human and Pseudomonas aeruginosa. Tncer showed unique biochemical properties. The optimum pH for Tncer activity is about pH5. From pH5to pH12, the ceramidase activity did not change too much. Tncer could hydrolyze various ceramidase from C6-ceramide to C24:1-ceramide. However the hydrolyzed ability to C6-ceramide is lower. The best temperature for its in vitro activity is about37℃. Tncer is a cation independent enzyme, Fe2+can inhibit its activity. The mRNA level of Tncer is highest in spermary and ovary of Tribolium castaneum. And the mRNA level of Tncer is increased with the maturation of ovary. These suggest that Tncer may paly roles in the oogenesis. Tncer is mainly located in the plasma membrane in High Five cells.
2. With the sequence assembly, we got the neutral ceramidase homologue (LsnCer) from the transcriptome database of Laodelphax striatellus. The sequence alignment showed that the sequence similarities between LsnCer and neutral ceramidase of Tribolium castaneum, Drosophila melanogaster, human, Pseudomonas aeruginosa and rice is50.62%,47.76%,37.11%,36.14%and36.84%respectively. The optimum pH for LsnCer is pH8. Little activity was detected in the strong acid and alkaline pH environment. It prefers medium-chain to long-chain ceramide as substrates. Short chain-ceramides such as C2-ceramide and super long chain-ceramide are resistant to LsnCer. The activity of LsnCer could be inhibited significantly by Zn2+The best temperature for its in vitro activity is also37℃. The mRNA level of LsnCer is highest in spermary and ovary of Laodelphax striatellus. It's a transmembrane protein and located in the plasma membrane in the High Five cells.
3. Rice stripe virus (RSV) can stimulate the expression of LsnCer. The ceramidase activity of LsnCer around pH8is also higher in RSV-infected Laodelphax striatellus. mRNA level of LsnCer as well as the neutral ceramidase activity were stimulated significantly after treated with insecticide4hours later.
4. We cloned neutral ceramidase1homologue (LsnSMase) from Laodelphax striatellus.323-342AA and355-374AA are two transmembrane domains. With the bioinformatic analysis and reported results, we presumed LsnSMase is the main sphingomyelinase in Laodelphax striatellus. The activity of LsnSMase is dependent on Mg2+, DTT can also increased the activity of LsnSMase. EDTA inhibited LsnSMase activity significantly. However, EGTA is better stimulator of LsnSMase than Mg2+. The co-operation of Mg2+and EGTA or Mg2+and EDTA or Mg2+and DTT could increase the LsnSMase activity sharply. mRNA level of LsnSMase is highest in ovary of Laodelphax striatellus.
In conclusion, we first identified biochemical properties of neutral ceramidase and neutral sphingomyelinase of agriculture pest. The function of these enzymes are also uncovered in our research.
引文
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