丽蝇蛹集金小蜂毒液性质及其寄生的生理学效应
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摘要
为了探明丽蝇蛹集金小蜂如何调控寄主免疫的机理，本论文就该蜂毒腺结构，毒液成份以及寄生对寄主血细胞数量和组成，血淋巴可溶性蛋白组份、含量，以及糖含量的变化作了初步的研究，结果如下：
未寄生的棕尾别麻蝇Boettcherisca peregrina蛹血淋巴中主要存在两种血细胞，即浆血细胞和颗粒细胞。棕尾别麻蝇蛹被丽蝇蛹集金小蜂Nasonia nitripennis寄生后1d浆血细胞的比例显著高于同期未寄生蛹，颗粒细胞比例也显著高于同期未寄生蛹。但寄生2～5d的蛹与同期未寄生蛹的颗粒细胞、浆血细胞占总细胞数的比例没有显著差异。数量方面，被寄生蛹的颗粒细胞和浆血细胞数量在寄生后1～5 d均显著高于同期未寄生蛹。寄生1 d后寄生蛹与未寄生蛹的血细胞总数(THC)间没有明显差异。寄生后2～5d，被寄生蛹的THC显著高于同期未寄生蛹，分别为同期为寄生蛹的3.2、1.8、2.1和2.3倍。
毒液器官的超微结构研究结果表明，毒囊包括一层外肌肉层，单层分泌细胞层和平行于内腔，贴近内膜的单层鳞状细胞层。内膜内陷到各个分泌细胞间，形成一个外面由管状细胞包裹的管腔。而毒腺则是由立方形分泌细胞形成的单细胞层，和平行于内腔，贴近内膜的单层鳞状细胞层共同构成。丽蝇蛹集金小蜂毒囊肌肉层是由纵向的肌肉组成的，相对较厚，毒囊没有分泌细胞，并受神经支配，寄生时毒液的注射是由肌肉层的收缩而造成的。丽蝇蛹金小蜂毒腺由单立方形分泌细胞层，和内部贴近内腔的鳞片状的细胞层共同构成的。分泌细胞核相对较大，并且有分散的染色质，常常可以看到分泌的毒液颗粒。围绕着分泌细胞的粗面内质网及高尔基体形成的小泡浓度充分表明了分泌器官具有活跃的分泌和运输功能，但在毒囊细胞中则很少可以看到，充分表明分泌功能主要在毒腺，而毒囊的主要功能则是贮藏，分泌功能则较弱。
利用PDQuest~(TM)二维电泳图像分析软件分析毒液的双向电泳图谱，可以识别到100多个毒液蛋白质点。从蛋白质的分子量分布范围来分析，分子量从2KDa到145KDa的区域均有蛋白质点，大于70KDa的蛋白质点很少，分子量在2KDa-40KDa蛋白质点较密。从蛋白质点的等电点分别范围来分析，偏于酸性端的蛋白质点明显多于碱性端的蛋白质点，大多数蛋白质点的等电点为中性或酸
性。
丽蝇蛹集金小蜂羽化后0．5．6d毒液含量整体呈逐渐上升的趋势，其中第6d
蛋白质浓度达到最高。而第7d蛋白质浓度则急剧下降，显著低于第6d的浓度。
寄生后 l－4d蛹血淋巴蛋白浓度相对稳定，在22．95－43．sling／ffil之间变化。
未寄生血淋巴蛋白浓度均极显著高于同期寄生蛹，分别是寄生蛹血淋巴蛋白浓度
的2．2、1．9、2．4和1．5倍。未寄生蛹血淋巴蛋白浓度1～4d逐渐下降；而寄生蛹
则与此不同，寄生后l—3d逐渐下降，到了4d则稍稍上升，保持在一个稳定的
水平。100 KDa的蛋白在棕尾别麻蝇蛹寄生初期含量较低，寄生后第 3 d寄生蛹
中此蛋白的含量明显升高，而到同期未寄生蛹中第4d此蛋白的含量才有所上升，
但远达不到被寄生蛹中的水平。贮藏蛋白在寄生蛹体内降解明显，同时，32 KDa
和 28 KDa蛋白在寄生 2 d后含量下降。
寄生后 1叫 d蛹血淋巴中可溶性糖含量较为稳定，而未寄生蛹血淋巴中可溶
性糖的含量在寄生后上升，寄生Zd后达到最高而后又下降。寄生蛹血淋巴中可
溶性糖含量在寄生后1～4d显著低于同期未寄生蛹；未寄生蛹血淋巴可溶性糖含
量在Zd后达到最高，而寄生蛹则在4d后达到最高。
In order to investigate the mechanism how to regulate the host's immunity by Nasonia nitripennis ,this article makes the preliminary study in the following topics: the structure of venom gland, change in constitution and count of hemocyte population, and the concentration of soluble proteins and soluble sugars after parasitization. The result is summarized as following:
There are two kinds of hemocyte in hemolymph of Boettcherisca peregrina pupae, plasmatoctes(PLs) and granular cells(GRs). In the first day after parasitization, the percentage of PLs is significantly higher in parasitized pupae than unparasitized ones, so as the percentage of GRs. But from the second day to the fifth day after parasitized, the percentage of GRs and PLs aren't significantly different between the parasitized pupaes and in unparasitized pupaes.
From the first day to the fifth day after parasitized, the count of GRs and PLs is significantly higher in parasitized pupaes than in unparasitized pupaes. After the first day parasitzed, total hemocyte count (THC) isn't significantly different between parasitized pupaes and unparasitized ones, and from the second day to the fifth day, it is significantly higher in parasitized pupaes than unparasitized ones, and THC of parasitized pupaes is 3.2, 1.8, 2.1 and 2.3 times than that of unparasitized pupaes.
The venom reservoir is consisted of an outer musular sheath, a single layer of secretory cells and an inner single layer of squamous cells adjacents to the intima which lines the lumen. The intima invaginates into each secretory cell and forms a duct which is surrounded by duct cells. And the gland filaments consist of a single layer of cuboidal secretory cells and an inner layer of squamous cells adjacent to the intima which lines the lumen. The reservoir apparatus are bordered distally by longitudinal muscles which form a relatively thick muscular sheath, and are innervated, the rapid injection of the venom may be facilitated by muscular contraction of the reservoir. In the gland filaments, the secretory cells of venom apparatus are cuboidal and form a continuous layer which is limited basically by a granular lamella. Nuclei of secretory cells are relatively large and contain scattered
chromatin. Concentration of sER and Golgi bodies around the secretory appartus indicate the possibility of active transport through the secretory appartus. But there can't be seen in the reservoir's cells. From the fact above, we can make a conclusion that the secretory function mainly acted by the gland filament, and the main function of reservoir is storing the venom.
From the utilization of the software PDQuest檛o exmaine the photo of venom's electrophoresis, there are 100 more protein dots can be deteced. The protein's weight are from 2 KDa to 145 KDa, the dots mainly scattered in the weight from 2 KDa to 40 KDa, only several dots above the 70 KDa. Using 2-DE we know that venom protein are mainly acid protein, and molecular weight is mainly between 2-40 KDa.
From the zero point fifth day to the sixth day, the concentration of venom is ascending, and there comes the highest concentration. But on the seventh day, the concentration is sharply droping, and high lower than the concentration of sixth day.
The concentration of protein is relatively stably in the hemolymph from the first day to the fourth day, changing between 22.95mg/ml and 43. 51mg/ml, it is is 3.2, 1.8, 2.1, 2.3 times higher in unparasitized pupaes than the in parasitized pupaes. The concentration of protein in the hemolymph from the unparasitized pupae is droping from the first day to the fourth day. The concentration of protein in the hemolymph from parasitized pupaes are droping from the first day to the third day, but on the fourth day , concentration are rising. The concentration of the protein of 100 KDa is lower in the hemolymph of former stage, the concentration is rising on the third day in the parasitized pupaes, but rising on the fourth day in the unparasitized pupaes, much lower than that in parasitized pupaes. The c
未寄生的棕尾别麻蝇Boettcherisca peregrina蛹血淋巴中主要存在两种血细胞，即浆血细胞和颗粒细胞。棕尾别麻蝇蛹被丽蝇蛹集金小蜂Nasonia nitripennis寄生后1d浆血细胞的比例显著高于同期未寄生蛹，颗粒细胞比例也显著高于同期未寄生蛹。但寄生2～5d的蛹与同期未寄生蛹的颗粒细胞、浆血细胞占总细胞数的比例没有显著差异。数量方面，被寄生蛹的颗粒细胞和浆血细胞数量在寄生后1～5 d均显著高于同期未寄生蛹。寄生1 d后寄生蛹与未寄生蛹的血细胞总数(THC)间没有明显差异。寄生后2～5d，被寄生蛹的THC显著高于同期未寄生蛹，分别为同期为寄生蛹的3.2、1.8、2.1和2.3倍。
毒液器官的超微结构研究结果表明，毒囊包括一层外肌肉层，单层分泌细胞层和平行于内腔，贴近内膜的单层鳞状细胞层。内膜内陷到各个分泌细胞间，形成一个外面由管状细胞包裹的管腔。而毒腺则是由立方形分泌细胞形成的单细胞层，和平行于内腔，贴近内膜的单层鳞状细胞层共同构成。丽蝇蛹集金小蜂毒囊肌肉层是由纵向的肌肉组成的，相对较厚，毒囊没有分泌细胞，并受神经支配，寄生时毒液的注射是由肌肉层的收缩而造成的。丽蝇蛹金小蜂毒腺由单立方形分泌细胞层，和内部贴近内腔的鳞片状的细胞层共同构成的。分泌细胞核相对较大，并且有分散的染色质，常常可以看到分泌的毒液颗粒。围绕着分泌细胞的粗面内质网及高尔基体形成的小泡浓度充分表明了分泌器官具有活跃的分泌和运输功能，但在毒囊细胞中则很少可以看到，充分表明分泌功能主要在毒腺，而毒囊的主要功能则是贮藏，分泌功能则较弱。
利用PDQuest~(TM)二维电泳图像分析软件分析毒液的双向电泳图谱，可以识别到100多个毒液蛋白质点。从蛋白质的分子量分布范围来分析，分子量从2KDa到145KDa的区域均有蛋白质点，大于70KDa的蛋白质点很少，分子量在2KDa-40KDa蛋白质点较密。从蛋白质点的等电点分别范围来分析，偏于酸性端的蛋白质点明显多于碱性端的蛋白质点，大多数蛋白质点的等电点为中性或酸
性。
丽蝇蛹集金小蜂羽化后0．5．6d毒液含量整体呈逐渐上升的趋势，其中第6d
蛋白质浓度达到最高。而第7d蛋白质浓度则急剧下降，显著低于第6d的浓度。
寄生后 l－4d蛹血淋巴蛋白浓度相对稳定，在22．95－43．sling／ffil之间变化。
未寄生血淋巴蛋白浓度均极显著高于同期寄生蛹，分别是寄生蛹血淋巴蛋白浓度
的2．2、1．9、2．4和1．5倍。未寄生蛹血淋巴蛋白浓度1～4d逐渐下降；而寄生蛹
则与此不同，寄生后l—3d逐渐下降，到了4d则稍稍上升，保持在一个稳定的
水平。100 KDa的蛋白在棕尾别麻蝇蛹寄生初期含量较低，寄生后第 3 d寄生蛹
中此蛋白的含量明显升高，而到同期未寄生蛹中第4d此蛋白的含量才有所上升，
但远达不到被寄生蛹中的水平。贮藏蛋白在寄生蛹体内降解明显，同时，32 KDa
和 28 KDa蛋白在寄生 2 d后含量下降。
寄生后 1叫 d蛹血淋巴中可溶性糖含量较为稳定，而未寄生蛹血淋巴中可溶
性糖的含量在寄生后上升，寄生Zd后达到最高而后又下降。寄生蛹血淋巴中可
溶性糖含量在寄生后1～4d显著低于同期未寄生蛹；未寄生蛹血淋巴可溶性糖含
量在Zd后达到最高，而寄生蛹则在4d后达到最高。
In order to investigate the mechanism how to regulate the host's immunity by Nasonia nitripennis ,this article makes the preliminary study in the following topics: the structure of venom gland, change in constitution and count of hemocyte population, and the concentration of soluble proteins and soluble sugars after parasitization. The result is summarized as following:
There are two kinds of hemocyte in hemolymph of Boettcherisca peregrina pupae, plasmatoctes(PLs) and granular cells(GRs). In the first day after parasitization, the percentage of PLs is significantly higher in parasitized pupae than unparasitized ones, so as the percentage of GRs. But from the second day to the fifth day after parasitized, the percentage of GRs and PLs aren't significantly different between the parasitized pupaes and in unparasitized pupaes.
From the first day to the fifth day after parasitized, the count of GRs and PLs is significantly higher in parasitized pupaes than in unparasitized pupaes. After the first day parasitzed, total hemocyte count (THC) isn't significantly different between parasitized pupaes and unparasitized ones, and from the second day to the fifth day, it is significantly higher in parasitized pupaes than unparasitized ones, and THC of parasitized pupaes is 3.2, 1.8, 2.1 and 2.3 times than that of unparasitized pupaes.
The venom reservoir is consisted of an outer musular sheath, a single layer of secretory cells and an inner single layer of squamous cells adjacents to the intima which lines the lumen. The intima invaginates into each secretory cell and forms a duct which is surrounded by duct cells. And the gland filaments consist of a single layer of cuboidal secretory cells and an inner layer of squamous cells adjacent to the intima which lines the lumen. The reservoir apparatus are bordered distally by longitudinal muscles which form a relatively thick muscular sheath, and are innervated, the rapid injection of the venom may be facilitated by muscular contraction of the reservoir. In the gland filaments, the secretory cells of venom apparatus are cuboidal and form a continuous layer which is limited basically by a granular lamella. Nuclei of secretory cells are relatively large and contain scattered
chromatin. Concentration of sER and Golgi bodies around the secretory appartus indicate the possibility of active transport through the secretory appartus. But there can't be seen in the reservoir's cells. From the fact above, we can make a conclusion that the secretory function mainly acted by the gland filament, and the main function of reservoir is storing the venom.
From the utilization of the software PDQuest檛o exmaine the photo of venom's electrophoresis, there are 100 more protein dots can be deteced. The protein's weight are from 2 KDa to 145 KDa, the dots mainly scattered in the weight from 2 KDa to 40 KDa, only several dots above the 70 KDa. Using 2-DE we know that venom protein are mainly acid protein, and molecular weight is mainly between 2-40 KDa.
From the zero point fifth day to the sixth day, the concentration of venom is ascending, and there comes the highest concentration. But on the seventh day, the concentration is sharply droping, and high lower than the concentration of sixth day.
The concentration of protein is relatively stably in the hemolymph from the first day to the fourth day, changing between 22.95mg/ml and 43. 51mg/ml, it is is 3.2, 1.8, 2.1, 2.3 times higher in unparasitized pupaes than the in parasitized pupaes. The concentration of protein in the hemolymph from the unparasitized pupae is droping from the first day to the fourth day. The concentration of protein in the hemolymph from parasitized pupaes are droping from the first day to the third day, but on the fourth day , concentration are rising. The concentration of the protein of 100 KDa is lower in the hemolymph of former stage, the concentration is rising on the third day in the parasitized pupaes, but rising on the fourth day in the unparasitized pupaes, much lower than that in parasitized pupaes. The c
引文
王善青等．野外释放丽蝇蛹集金小蜂对家蝇及其它蝇种控制效果观察．中国媒介生物学及控制杂志．1993，4(1)：14-17
黄大卫．中国丽金小蜂属Lamprotatus Westwood(膜翅目：金小蜂科：柄腹金小蜂亚科)．动物分类学报．1991，16(2)：214-223
黄大卫．蝇蛹金小蜂的行为学研究．动物学集刊．1987，5：177—186
蔡峻．蝶蛹金小蜂调控寄主免疫机理的初步研究．浙江大学博士学位论文．2000
薛瑞德等．丽蝇蛹集金小蜂的微生态环境及其对个体发育的影响．生物学杂志．1987，5：10—14
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Jones D, et al.. Hemolymph concentrations of host ecdysteroids are strongly suppressed in precocious prepupae of Trichoplusia ni parasitized and pseudoparasitized by Chelonus near curvimaculatus. Archs. Insect Biochem. Physiol., 1992, 21: 155～165.
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95～106.
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Jones, D. Biochemical interaction between Chelonine wasps and their Lepidopteran host: After a decade of research-The parasite is in control., Insect. Biochem. Molec. Biol. 1996, 26: 981～996.
Jones, D., Wache, S. Preultimate 4th/5th Trichoplusia ni naturally-injected with venom/calyx fluid from Chelonus curvimaculatus precociously metamorphose, rather than obey the metamorphic size threshold that would normally compel molting to a 5th/6th instar, J. Insect Physiol. 1998, 44: 755～765.
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Lawrence P O and Akin D. Virus-like particles from the poison glands of the parasitic wasp Biosteres longicandatus (Hymenoptera: Braconidae). Can. Zool., 1990, 68: 539～546.
Li, X and Webb, B A. Apparent functional role for a cysteine-rich polydnavirus proteins in suppression of the insect cellular immune response. J. Virol., 1994,68: 7482-7489.
Marris G C, et al.. The role of Pimpla hypochondriaca venom in the suppression of pupal Nocuid host immunity. Entomol. Exp. Appl., 1999, 93: 291～298.
O'Farrell P H.High resolution Two-dimentional electrophoresis of proteins. J Bid Chem, 1975, 250:4007-4021
Okuda T, Kadono, Okuda K. Perilitus coccinellae teratocyte polypeptide: Evidence for production of a teratocyte-specific 540KDa protein. J. Insect Physiol., 1995,41:819-825
Park D.S., Shin S.W., Kim M.G. et al. Isolation and Characterization of the cDNA Encoding the Prophenoloxidase of Fall Webworm, Hyphantria cunea Insect Biochem. Mol. Biol., 1997,27:983～992
Parkinson N M, Weaver R J. Noxious components of venom from the pupa-specific parasitoid Pimpla hypochondriaca. J. Invert. Pathol. , 1999, 73: 74～83.
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黄大卫．中国丽金小蜂属Lamprotatus Westwood(膜翅目：金小蜂科：柄腹金小蜂亚科)．动物分类学报．1991，16(2)：214-223
黄大卫．蝇蛹金小蜂的行为学研究．动物学集刊．1987，5：177—186
蔡峻．蝶蛹金小蜂调控寄主免疫机理的初步研究．浙江大学博士学位论文．2000
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Jones D, et al.. Isomeric and quaternary properties of homogenous 33KDa protein from the venom of Chelonus near curvimaculatus. Archs. Insect Biochem. Physiol., 1994, 26: 83～95.
Jones D, Leluk J. Venom proteins of endoparasitic wasp Chelonus near curvimaculatus: characterization of the major components. Archs. Insect Biochem. Physiol., 1990, 13:
95～106.
Jones D, Wache S. Preultimate 4th/5th instar Trichoplusia ni naturally-injected with venom/calyx fluid from Chelonus curvimaculatus precociously metamorphose, rather than obey the metamorphic size threshold that would normally compel molting to a 5th/6th instar, J. Insect Physiol., 1998, 44: 755～765.
Jones, D. Biochemical interaction between Chelonine wasps and their Lepidopteran host: After a decade of research-The parasite is in control., Insect. Biochem. Molec. Biol. 1996, 26: 981～996.
Jones, D., Wache, S. Preultimate 4th/5th Trichoplusia ni naturally-injected with venom/calyx fluid from Chelonus curvimaculatus precociously metamorphose, rather than obey the metamorphic size threshold that would normally compel molting to a 5th/6th instar, J. Insect Physiol. 1998, 44: 755～765.
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Junnikkala, E. Testis development in Pieris brassicae parasitized by Apanteles glomeratus., Entomol. Exp. Appl., 1985, 37: 283.
Kamita, A.M., Majima, K., Maeda, S. Identification and characterization of the p35 gene of Bombyx mori nuclear virus that prevents virus-induced apoposis. J. Virol., 1993,67: 455～463.
Kitano H. Effect of venom of the gregarious parasitoid Apanteles glomeratus on its hemocytic encapsulation by the host, Pieris. J. Invert. Path., 1982, 40:61～67
Kitano H, 1986. The role of Apanteles glomeratus venom in the defensive response of its host Pieris rapae crucivora. J. Insect Physiol., 32: 369～375.
Kitano, H., Wago, H., Arakawa, T. Possible role of teratocytes of the gregarious parasitoid Cotesia(=Apanteles) glomerata in the suppression of phenoloxidase activity in the larval host, Pieris rapae crucivora. Archives of Insect Biochemistry and Physiology, 1990, 13:177-185
Krell, P.J., Summers, M.D., Vinson, S.B. Virus with a multipartite superhelical DNA genome from the Ichneumonid parasitoid Campoletis sonoransis. J. Virol., 1982, 43:859～870.
Krishnan A, et al.. Isolation, cloning and characterization of new chitinase stored in active form in chitin-lined venom reservoir, J. Biol. Chem., 1994, 269:20971～20976.
Lawrence P O and Akin D. Virus-like particles from the poison glands of the parasitic wasp Biosteres longicandatus (Hymenoptera: Braconidae). Can. Zool., 1990, 68: 539～546.
Li, X and Webb, B A. Apparent functional role for a cysteine-rich polydnavirus proteins in suppression of the insect cellular immune response. J. Virol., 1994,68: 7482-7489.
Marris G C, et al.. The role of Pimpla hypochondriaca venom in the suppression of pupal Nocuid host immunity. Entomol. Exp. Appl., 1999, 93: 291～298.
O'Farrell P H.High resolution Two-dimentional electrophoresis of proteins. J Bid Chem, 1975, 250:4007-4021
Okuda T, Kadono, Okuda K. Perilitus coccinellae teratocyte polypeptide: Evidence for production of a teratocyte-specific 540KDa protein. J. Insect Physiol., 1995,41:819-825
Park D.S., Shin S.W., Kim M.G. et al. Isolation and Characterization of the cDNA Encoding the Prophenoloxidase of Fall Webworm, Hyphantria cunea Insect Biochem. Mol. Biol., 1997,27:983～992
Parkinson N M, Weaver R J. Noxious components of venom from the pupa-specific parasitoid Pimpla hypochondriaca. J. Invert. Pathol. , 1999, 73: 74～83.
Parkinson N, et al.. A new form of arthropod phenoloxidase is abundant in venom of the parasitoid wasp Pimpla hypochondriaca. Insect Biochem. Mol. Biol., 2001, 31: 57～63.
Pennacchio, F., Falabella, P., Sordetti, R., et al. Prothoracic gland inactivation in Heliothis virescens Larvae parasitized by Cardiochiles nigriceps Viereck. J. Insect Phsiol., 1998, 44:845～857.
Pennacchio, F., Vison, S.B., Tremblay, E. Growth and development of Cardiochiles nigriceps Vieteck larvae and their synchronization with some changes of the hemolymph composition of their host, Heliothis virescens. Arch. Insect Biochem. Phsiol., 1993,24: 65～77.
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