半闭弯尾姬蜂调控寄主小菜蛾的生理机制研究
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摘要
本文对十字花科蔬菜重要害虫一小菜蛾Plutella xylostella幼虫的优势种寄生蜂半闭弯尾姬蜂Diadegma semiclausum对寄主的生理调控机制进行了研究，取得以下结果：
一、半闭弯尾姬蜂寄生对寄主免疫系统的影响寄主小菜蛾幼虫被寄生后，其造血器官内原血细胞结构受到一定程度的破坏，是造成血细胞数量显著下降的原因之一；但是血腔内成熟血细胞并不出现坏死或加速凋亡的现象。被寄生的寄主幼虫的血细胞总量在寄生后12 h呈现出相对于对照的显著低水平，且这种水平一直维持至寄主幼虫死亡。此外，寄生后1 h的寄主幼虫血细胞体外培养30 min后，不能延展，所有的细胞均呈现圆饼状，不具有包囊能力，黑化作用受到抑制，但噬菌能力并未受到影响；而寄生后24 h的寄主幼虫血细胞虽然已经能够进行正常的延展，恢复部分包囊能力，黑化作用的抑制程度减轻，但是其噬菌能力却显著降低。被半闭弯尾姬蜂寄生后的寄主体液免疫能力降低，如酚氧化酶酶原的激活受抑制、血淋巴抗菌效能降低等。
二、半闭弯尾姬蜂寄生对寄主解毒酶系统和保护酶的影响
半闭弯尾姬蜂寄生对寄主幼虫的解毒酶系统产生一定程度的影响，主要结果为：1)寄生能够显著提高寄主幼虫体内谷胱甘肽-s-转移酶的活性；2)多功能氧化酶和羧酸酯酶的酶活力不受影响；3)半闭弯尾姬蜂寄生显著降低乙酰胆碱酯酶的活性。
另外，寄生后小菜蛾幼虫的保护酶活性也发生一定的变化：1)过氧化物歧化酶(Superoxidedismutase，SOD)的活性在寄生后12 h显著降低至对照的50%左右，但是在此后12 h该酶活性恢复至正常水平；2)过氧化氢酶(Catalase，CAT)的活力不受半闭弯尾姬蜂寄生的影响；3)过氧化物酶(Peroxisome，POD)活力变化显著，寄生后1 h即与对照呈显著差异。
三、半闭弯尾姬蜂寄生对寄主营养代谢的影响
在小菜蛾-半闭弯尾姬蜂系统中，寄主幼虫血淋巴内海藻糖浓度升高的同时脂肪体内糖原积累受到抑制；在小菜蛾-菜蛾盘绒茧蜂系统中，寄主幼虫血淋巴内海藻糖浓度也显著升高，而且这种现象的产生与寄主幼虫体内脂肪体糖原的加速降解和脂滴转移有关。在两个系统中，虽然寄生后幼虫的糖原磷酸化酶及海藻糖酶的活力发生一定程度的变化，但是这两种酶的变化并不直接导致海藻糖浓度的升高。
寄生后小菜蛾幼虫的血淋巴内蛋白总量发生显著变化，同时中肠内各类消化酶之间的协调性被打破。总胰蛋白酶、弱碱性类胰蛋白酶、强碱性类胰蛋白酶活力均发生不同程度的改变，且这些酶类对酸碱环境的适应性也发生了一定程度上的变化。
在脂类代谢方面，半闭弯尾姬蜂寄生对小菜蛾脂肪体重量的影响主要发生在幼虫后期及预蛹期。在4龄后期，对照幼虫脂肪体开始不断积累营养物质，脂肪体重量逐渐增大；而寄生后幼虫体内的脂肪体总量却呈现下降的趋势。半闭弯尾姬蜂对寄主脂肪体的利用分寄主幼虫期和寄主预蛹期两个阶段进行：寄主幼虫期阶段，幼蜂的总脂滴含量与小菜蛾幼虫的血淋巴总脂滴含量显著相关(P=0．2976H+4．9877，r~2=0．9824；P代表幼蜂总脂滴含量，H代表小菜蛾幼虫血淋巴的总脂滴含量)；寄主预蛹阶段，幼蜂的总脂滴含量与寄主脂肪体的总脂滴含量显著相关(P=476．61-56．705 Ln(F)，r~2=0．9887；P代表幼蜂总脂滴含量，F代表小菜蛾预蛹脂肪体中总脂滴含量)，但与血淋巴内总脂滴含量的相关性并不显著。对小菜蛾幼虫及预蛹的血淋巴及脂肪体和半闭弯尾姬蜂幼虫体内所含有机质进行抽提及GC-MS分析后结果发现，寄生及未寄生小菜蛾幼虫的血淋巴、脂肪体及半闭弯尾姬蜂幼蜂体内的有机质组分相似；但是，寄生与未寄生小菜蛾预蛹的脂肪体内有机质组分却存在很大差异。
四、半闭弯尾姬蜂寄生对寄主取食、生长及代谢效率的影响
被半闭弯尾姬蜂寄生后的小菜蛾幼虫的代谢率明显提高，且显著高于被菜蛾盘绒茧蜂寄生后寄主的代谢率。在小菜蛾-半闭弯尾姬蜂系统中，寄主的取食、生长及其他营养代谢相关参数受到一定程度的影响，但这些变化的显著性相对弱于小菜蛾-菜蛾盘绒茧蜂系统。被菜蛾盘绒茧蜂寄生后小菜蛾幼虫的相对生长率、取食量、排泄量显著降低，同化作用被显著抑制。寄主的呼吸作用，在寄生后的24 h内受抑制，但针对于整个寄生期而言，受抑制现象并不显著；在寄主和寄生蜂共生期间，寄主的消化食物转化率、摄入食物转化率降低，而近似消化率则上升。
五、半闭弯尾姬蜂寄生对寄主部分内部器官变化的影响
半闭弯尾姬蜂寄生后寄主幼虫内部器官的结构变化不明显。但是，菜蛾盘绒茧蜂寄生后的小菜蛾幼虫部分内部器官结构发生显著变化，如丝腺上的功能细胞在未完全转化成为功能型细胞之前发生凋亡，即分泌细胞丧失分泌功能。寄主脑内神经细胞的超微结构也发生一些变化，如核畸形或肿胀，电子密度发生变化，胞膜消失，内容物降解，染色质溶解，核仁呈降解状；细胞间隙变大，胞间质稀薄，胞间连接(如细胞桥粒等)消失；突触囊泡内内容物的泡状结构被破坏，数量减少，大小不均一；寄生后的神经分泌类物质较未寄生的幼虫明显减少等。
六、半闭弯尾姬蜂个体发育及寄生因子研究
半闭弯尾姬蜂蜂卵属于膜翅目型，卵表面无附属物。幼虫龄期分为4龄。蛹期分为两个阶段：第一阶段形似于5龄幼虫，但其体表会出现一些弱的突起；第二阶段形态呈裸蛹，其蛹体表各部位的着色顺序依次为：复眼-单眼-下颚-胸-腹-触角。
首次发现半闭弯尾姬蜂的寄生因子除了毒液外，还携有另一个重要的寄生因子—多分DNA病毒(Diadegma semiclausum ichnivirus，DsIVs)。每个毒液囊的蛋白含量为1．016μg左右，大部分毒液蛋白的分子量集中在35-220 kDa范围内。DsIVs粒子是典型的姬蜂病毒，病毒粒子在萼细胞内获得第一层外膜，而在以出芽的方式进入萼区内腔时获得第二层外膜；成熟的病毒粒子呈圆柱状，横截面直径为130-150 nm而长度在300-400 nm之间，中间为高浓度的核质。
半闭弯尾姬蜂卵巢由12根卵巢小管组成，属多滋式。超微结构表明卵巢管内配子囊中含多个细胞，但只有一个细胞可发育成为卵细胞，而其他细胞则分化为滋养细胞为卵细胞的发育提供营养，在卵细胞接近成熟的同时逐步消亡。刚羽化的雌蜂体内萼区为空，不含有成熟的卵；在羽化后2-3 d，萼区即充满了成熟的卵细胞。成熟卵细胞进入萼区后，被含DsIVs粒子的萼液包裹，但DsIVs粒子并不会附着到蜂卵绒毛膜表面。
Diadegma semiclausum (Hell(?)n) (Hymenoptera:Ichneumonidae) is an important naturalenemy of the diamondback moth,Plutella xylostella L.(Lepidoptera:Plutellidae) in manyregions of the world.The physiological effects of parasitism by this endoparasitoid on itshost were studied in this dissertation.The major results are summarized as follows:
1 Immune change of P.xylostella larvae parasitized by D.semiclausum
Necrosis ofprohemocytes in hematopoietic organ was observed while cell death wasabsent in the mature hemocytes in the parasitized larvae,which was related to the declinedtotal hemocyte count per microliter (THC).THC in the host hemolymph declined sharply by12 h post-parasitization and then remained at a low level.When hemocvtes of the parasitizedlarvae were cultured in vitro,encapsulation ability was suppressed coincidently with theinhibited spreading ability and depressed melanization;but phagocytosis ability of thehemocytes was not affected.After 24 hours,spreading,encapsulation and melanization ofthe hemocytes in the parasitized host larvae could be normally occurred;however.phagocytosis ability was significantly inhibited.In the aspect of humoral immune system inthe parasitized host larvae,activation of the prophenoloxidae was inhibited,and thebacteriostatic ability of hemolymph was decreased.
2 Effect of parasitism by D.semiclausum on the activities of detoxification system andprotective enzymes of its host larva
The detoxification enzymes of host larva were altered by parasitism,resulting in (1) theactivity of glutathione -s- transferase in the host larvae was increased after parasitization;(2)activities of mixed-functionoxidases or carboxylesterase were not significantly affected;(3)activity of acetylcholinesterase was decreased.
The protective enzymes of host larva were also altered by parasitism,and the resultswere as follows:(1) activity of superoxide dismutase was decreased to the half level of thecontrol,however,this enzyme was reactivated after 12 hours post parasitism;(2) activity ofcatalase was not affected;and (3) activity of peroxisome was significantly altered.
3 Effect of parasitism by D.semiclausum on the host larval nutrient metabolism
Hypertrehalosemia was observed in P.xylostella larvae parasitized by either C.vestalisor D.semiclausum.Results of our experiments showed that it was caused by theaccelerated degradation of glycogen in the fat body and probably by lipid mobilization inthe larvae parasitized by C.vestalis.However,increased trehalose in the larvae parasitizedby D.semiclausum accompanied by decreased accumulation of glycogen in the fat bodiesand unchanged lipid concentration.Additionally,it is demonstrated that neither glycogenphosphorylase nor trehalase was responsible for elevated trehalose in the parasitizedlepidopteran larvae.
Moreover,protein content in the hemolymph was significantly affected by parasitism,and the balance among the proteinases in the midguts was disrupted.Activities oftrypsinase/BApNAase/TAMEase in the parasitized larvae were drastically changed;andfurthermore,sensitivities of these three enzymes to pH were also altered.
Effect of parasitism by D.semiclausum on the host lipid metabolism was mainlyhappened in the late host larval and prepupal stages.The late 4th instar larvae began toaccmulate substance in fat body,and the weight of fat body was gradually increased;however,the fat body weight of the parasitized larvae in the same stages showed adecreasing trend.Change of the amount of total lipid in D.semiclausum was significantlycorrelated with that of host larval haemolymph (P = 0.2976 H + 4.9877,r~2= 0.9824;P:total lipid in D.semiclausum larva,H:total lipid in P.xylostella larval hemolymph) andpre-pupal fat body (P = 476.61-56.705Ln(F),r~2=0.9887;P:total lipid in D.semiclausumlarva,H:total lipid in P.xylostella prepupal fatbody).Profiles of organic compound fromparasitized/non-parasitized larval haemolymph/fat body,and parasitoid larva were similarwith that from parasitzed pre-pupal fat bodies;however,that from non-parasitizedpre-pupal fat bodies was different.
4 Parasitism-induced effects on metabolic efficiency and host growth in P.xylostellalarvae parasitized by D.semiclausum
Parasitization by C.vestalis resulted in significant decrease in the rates of growth,feeding,excretion,assimilation,and respiration,but the final dry rate of respiration at theend of last larval stadium was elevated.The ECI (.percent efficiency of conversion of ingested food to body substance) and ECD (percent efficiency of conversion of digestedfood to body substance) were also reduced as the result of parasitization,but digestibilitywas increased.All these parameters in the larvae parasitized by D.semiclausum at 24 hpost-parasitization were changed much less significantly.In spite of the similarities of theparasitism-induced effects caused by these endoparasitoids,the final metabolic rate,that is,the rate of intake of nutrients required to compensate for metabolism,was much lower inthe larvae parasitized by C.vestalis than that in the larvae parasitized by D.semiclausum.
5 Changes in some organ of the host larva parasitized by D.semiclausum
After parasitized by C.vestalis,structures of some organs in host larvae were altered,forexample,cell death was happened in the cells,which could be developed into secretory cells,of the silk gland before the cellular differentiation was completed.Some ultrastrcuturalchanges in neural cells in the brain were presented,i.e.,tumefied/abnormal karyon,alteredelectronica density,disappeard cellular membrane,cytoplasmic degradation.Futhermore,gap between cells became larger,cellular conjunction disappeared,synaptic vesicles weredisrupted,and accumulation of the neurosecretion was decreased.However,changesmentioned above were not so significant in the P.xylostella larvae parasitized by D.semiclausum.
6 Individual development of D.semiclausum:oogenesis with associated polydnavirus,external morphology and development of immature stages
The egg of D.semiclausum is hymenopteriform,lacking appendages.The larvae appearto be hypermetamorphic with four instars.The pupal period has two distinct phases:(a)phase I:appears larval-like with remarkable groups of urate cells;(b) phase II:exarate pupalappearance with a pigmentation sequence of eyes,ocelli,ma ndibles,thorax,abdomen,andantennae.
D.semiclausum female was found to carry a kind of PDVs (DsIV) as its majorparasitoid-associated factors beside venom for the first time.Protein content,most of whichwas ranged from 35 kDa to 220 kDa.in each venom reservior was 1.016μg.DslVs weretypical ichneumonid polydnaviruses with the following characteristics:( 1 ) the virus particles were assembled in the nuclei of calyx cells where they acquire an initial (inner) membrane,then migrate through the cytoplasm and budd out into the lumen of the ovary,and at thistime they acquire a second envelope (outer membrane);(2) the particle,which had adiameter of 130-150 nm with a length of 300-400 nm,has a genome comprising severalDNA segments.
Ovary of female D.semiclausum had 12 orarioles,and belonged to the polytrophic type.Ultrastructural observations demonstrated that several cells existed in a gametangium,andone of the central cells of the cluster differentiated into an oocyte while the remainingbecame nurse cells.Germ cells continued to develop in the vitellarium until mature;howeverthe nurse cells were in gradual degradation simultaneously.The calyx region was emptywhen the adult was just emerged;and after 2-3 days,mature eggs began to enter the calyxregion and surrounded by calyx fluid but without DsIVs attached.
一、半闭弯尾姬蜂寄生对寄主免疫系统的影响寄主小菜蛾幼虫被寄生后，其造血器官内原血细胞结构受到一定程度的破坏，是造成血细胞数量显著下降的原因之一；但是血腔内成熟血细胞并不出现坏死或加速凋亡的现象。被寄生的寄主幼虫的血细胞总量在寄生后12 h呈现出相对于对照的显著低水平，且这种水平一直维持至寄主幼虫死亡。此外，寄生后1 h的寄主幼虫血细胞体外培养30 min后，不能延展，所有的细胞均呈现圆饼状，不具有包囊能力，黑化作用受到抑制，但噬菌能力并未受到影响；而寄生后24 h的寄主幼虫血细胞虽然已经能够进行正常的延展，恢复部分包囊能力，黑化作用的抑制程度减轻，但是其噬菌能力却显著降低。被半闭弯尾姬蜂寄生后的寄主体液免疫能力降低，如酚氧化酶酶原的激活受抑制、血淋巴抗菌效能降低等。
二、半闭弯尾姬蜂寄生对寄主解毒酶系统和保护酶的影响
半闭弯尾姬蜂寄生对寄主幼虫的解毒酶系统产生一定程度的影响，主要结果为：1)寄生能够显著提高寄主幼虫体内谷胱甘肽-s-转移酶的活性；2)多功能氧化酶和羧酸酯酶的酶活力不受影响；3)半闭弯尾姬蜂寄生显著降低乙酰胆碱酯酶的活性。
另外，寄生后小菜蛾幼虫的保护酶活性也发生一定的变化：1)过氧化物歧化酶(Superoxidedismutase，SOD)的活性在寄生后12 h显著降低至对照的50%左右，但是在此后12 h该酶活性恢复至正常水平；2)过氧化氢酶(Catalase，CAT)的活力不受半闭弯尾姬蜂寄生的影响；3)过氧化物酶(Peroxisome，POD)活力变化显著，寄生后1 h即与对照呈显著差异。
三、半闭弯尾姬蜂寄生对寄主营养代谢的影响
在小菜蛾-半闭弯尾姬蜂系统中，寄主幼虫血淋巴内海藻糖浓度升高的同时脂肪体内糖原积累受到抑制；在小菜蛾-菜蛾盘绒茧蜂系统中，寄主幼虫血淋巴内海藻糖浓度也显著升高，而且这种现象的产生与寄主幼虫体内脂肪体糖原的加速降解和脂滴转移有关。在两个系统中，虽然寄生后幼虫的糖原磷酸化酶及海藻糖酶的活力发生一定程度的变化，但是这两种酶的变化并不直接导致海藻糖浓度的升高。
寄生后小菜蛾幼虫的血淋巴内蛋白总量发生显著变化，同时中肠内各类消化酶之间的协调性被打破。总胰蛋白酶、弱碱性类胰蛋白酶、强碱性类胰蛋白酶活力均发生不同程度的改变，且这些酶类对酸碱环境的适应性也发生了一定程度上的变化。
在脂类代谢方面，半闭弯尾姬蜂寄生对小菜蛾脂肪体重量的影响主要发生在幼虫后期及预蛹期。在4龄后期，对照幼虫脂肪体开始不断积累营养物质，脂肪体重量逐渐增大；而寄生后幼虫体内的脂肪体总量却呈现下降的趋势。半闭弯尾姬蜂对寄主脂肪体的利用分寄主幼虫期和寄主预蛹期两个阶段进行：寄主幼虫期阶段，幼蜂的总脂滴含量与小菜蛾幼虫的血淋巴总脂滴含量显著相关(P=0．2976H+4．9877，r~2=0．9824；P代表幼蜂总脂滴含量，H代表小菜蛾幼虫血淋巴的总脂滴含量)；寄主预蛹阶段，幼蜂的总脂滴含量与寄主脂肪体的总脂滴含量显著相关(P=476．61-56．705 Ln(F)，r~2=0．9887；P代表幼蜂总脂滴含量，F代表小菜蛾预蛹脂肪体中总脂滴含量)，但与血淋巴内总脂滴含量的相关性并不显著。对小菜蛾幼虫及预蛹的血淋巴及脂肪体和半闭弯尾姬蜂幼虫体内所含有机质进行抽提及GC-MS分析后结果发现，寄生及未寄生小菜蛾幼虫的血淋巴、脂肪体及半闭弯尾姬蜂幼蜂体内的有机质组分相似；但是，寄生与未寄生小菜蛾预蛹的脂肪体内有机质组分却存在很大差异。
四、半闭弯尾姬蜂寄生对寄主取食、生长及代谢效率的影响
被半闭弯尾姬蜂寄生后的小菜蛾幼虫的代谢率明显提高，且显著高于被菜蛾盘绒茧蜂寄生后寄主的代谢率。在小菜蛾-半闭弯尾姬蜂系统中，寄主的取食、生长及其他营养代谢相关参数受到一定程度的影响，但这些变化的显著性相对弱于小菜蛾-菜蛾盘绒茧蜂系统。被菜蛾盘绒茧蜂寄生后小菜蛾幼虫的相对生长率、取食量、排泄量显著降低，同化作用被显著抑制。寄主的呼吸作用，在寄生后的24 h内受抑制，但针对于整个寄生期而言，受抑制现象并不显著；在寄主和寄生蜂共生期间，寄主的消化食物转化率、摄入食物转化率降低，而近似消化率则上升。
五、半闭弯尾姬蜂寄生对寄主部分内部器官变化的影响
半闭弯尾姬蜂寄生后寄主幼虫内部器官的结构变化不明显。但是，菜蛾盘绒茧蜂寄生后的小菜蛾幼虫部分内部器官结构发生显著变化，如丝腺上的功能细胞在未完全转化成为功能型细胞之前发生凋亡，即分泌细胞丧失分泌功能。寄主脑内神经细胞的超微结构也发生一些变化，如核畸形或肿胀，电子密度发生变化，胞膜消失，内容物降解，染色质溶解，核仁呈降解状；细胞间隙变大，胞间质稀薄，胞间连接(如细胞桥粒等)消失；突触囊泡内内容物的泡状结构被破坏，数量减少，大小不均一；寄生后的神经分泌类物质较未寄生的幼虫明显减少等。
六、半闭弯尾姬蜂个体发育及寄生因子研究
半闭弯尾姬蜂蜂卵属于膜翅目型，卵表面无附属物。幼虫龄期分为4龄。蛹期分为两个阶段：第一阶段形似于5龄幼虫，但其体表会出现一些弱的突起；第二阶段形态呈裸蛹，其蛹体表各部位的着色顺序依次为：复眼-单眼-下颚-胸-腹-触角。
首次发现半闭弯尾姬蜂的寄生因子除了毒液外，还携有另一个重要的寄生因子—多分DNA病毒(Diadegma semiclausum ichnivirus，DsIVs)。每个毒液囊的蛋白含量为1．016μg左右，大部分毒液蛋白的分子量集中在35-220 kDa范围内。DsIVs粒子是典型的姬蜂病毒，病毒粒子在萼细胞内获得第一层外膜，而在以出芽的方式进入萼区内腔时获得第二层外膜；成熟的病毒粒子呈圆柱状，横截面直径为130-150 nm而长度在300-400 nm之间，中间为高浓度的核质。
半闭弯尾姬蜂卵巢由12根卵巢小管组成，属多滋式。超微结构表明卵巢管内配子囊中含多个细胞，但只有一个细胞可发育成为卵细胞，而其他细胞则分化为滋养细胞为卵细胞的发育提供营养，在卵细胞接近成熟的同时逐步消亡。刚羽化的雌蜂体内萼区为空，不含有成熟的卵；在羽化后2-3 d，萼区即充满了成熟的卵细胞。成熟卵细胞进入萼区后，被含DsIVs粒子的萼液包裹，但DsIVs粒子并不会附着到蜂卵绒毛膜表面。
Diadegma semiclausum (Hell(?)n) (Hymenoptera:Ichneumonidae) is an important naturalenemy of the diamondback moth,Plutella xylostella L.(Lepidoptera:Plutellidae) in manyregions of the world.The physiological effects of parasitism by this endoparasitoid on itshost were studied in this dissertation.The major results are summarized as follows:
1 Immune change of P.xylostella larvae parasitized by D.semiclausum
Necrosis ofprohemocytes in hematopoietic organ was observed while cell death wasabsent in the mature hemocytes in the parasitized larvae,which was related to the declinedtotal hemocyte count per microliter (THC).THC in the host hemolymph declined sharply by12 h post-parasitization and then remained at a low level.When hemocvtes of the parasitizedlarvae were cultured in vitro,encapsulation ability was suppressed coincidently with theinhibited spreading ability and depressed melanization;but phagocytosis ability of thehemocytes was not affected.After 24 hours,spreading,encapsulation and melanization ofthe hemocytes in the parasitized host larvae could be normally occurred;however.phagocytosis ability was significantly inhibited.In the aspect of humoral immune system inthe parasitized host larvae,activation of the prophenoloxidae was inhibited,and thebacteriostatic ability of hemolymph was decreased.
2 Effect of parasitism by D.semiclausum on the activities of detoxification system andprotective enzymes of its host larva
The detoxification enzymes of host larva were altered by parasitism,resulting in (1) theactivity of glutathione -s- transferase in the host larvae was increased after parasitization;(2)activities of mixed-functionoxidases or carboxylesterase were not significantly affected;(3)activity of acetylcholinesterase was decreased.
The protective enzymes of host larva were also altered by parasitism,and the resultswere as follows:(1) activity of superoxide dismutase was decreased to the half level of thecontrol,however,this enzyme was reactivated after 12 hours post parasitism;(2) activity ofcatalase was not affected;and (3) activity of peroxisome was significantly altered.
3 Effect of parasitism by D.semiclausum on the host larval nutrient metabolism
Hypertrehalosemia was observed in P.xylostella larvae parasitized by either C.vestalisor D.semiclausum.Results of our experiments showed that it was caused by theaccelerated degradation of glycogen in the fat body and probably by lipid mobilization inthe larvae parasitized by C.vestalis.However,increased trehalose in the larvae parasitizedby D.semiclausum accompanied by decreased accumulation of glycogen in the fat bodiesand unchanged lipid concentration.Additionally,it is demonstrated that neither glycogenphosphorylase nor trehalase was responsible for elevated trehalose in the parasitizedlepidopteran larvae.
Moreover,protein content in the hemolymph was significantly affected by parasitism,and the balance among the proteinases in the midguts was disrupted.Activities oftrypsinase/BApNAase/TAMEase in the parasitized larvae were drastically changed;andfurthermore,sensitivities of these three enzymes to pH were also altered.
Effect of parasitism by D.semiclausum on the host lipid metabolism was mainlyhappened in the late host larval and prepupal stages.The late 4th instar larvae began toaccmulate substance in fat body,and the weight of fat body was gradually increased;however,the fat body weight of the parasitized larvae in the same stages showed adecreasing trend.Change of the amount of total lipid in D.semiclausum was significantlycorrelated with that of host larval haemolymph (P = 0.2976 H + 4.9877,r~2= 0.9824;P:total lipid in D.semiclausum larva,H:total lipid in P.xylostella larval hemolymph) andpre-pupal fat body (P = 476.61-56.705Ln(F),r~2=0.9887;P:total lipid in D.semiclausumlarva,H:total lipid in P.xylostella prepupal fatbody).Profiles of organic compound fromparasitized/non-parasitized larval haemolymph/fat body,and parasitoid larva were similarwith that from parasitzed pre-pupal fat bodies;however,that from non-parasitizedpre-pupal fat bodies was different.
4 Parasitism-induced effects on metabolic efficiency and host growth in P.xylostellalarvae parasitized by D.semiclausum
Parasitization by C.vestalis resulted in significant decrease in the rates of growth,feeding,excretion,assimilation,and respiration,but the final dry rate of respiration at theend of last larval stadium was elevated.The ECI (.percent efficiency of conversion of ingested food to body substance) and ECD (percent efficiency of conversion of digestedfood to body substance) were also reduced as the result of parasitization,but digestibilitywas increased.All these parameters in the larvae parasitized by D.semiclausum at 24 hpost-parasitization were changed much less significantly.In spite of the similarities of theparasitism-induced effects caused by these endoparasitoids,the final metabolic rate,that is,the rate of intake of nutrients required to compensate for metabolism,was much lower inthe larvae parasitized by C.vestalis than that in the larvae parasitized by D.semiclausum.
5 Changes in some organ of the host larva parasitized by D.semiclausum
After parasitized by C.vestalis,structures of some organs in host larvae were altered,forexample,cell death was happened in the cells,which could be developed into secretory cells,of the silk gland before the cellular differentiation was completed.Some ultrastrcuturalchanges in neural cells in the brain were presented,i.e.,tumefied/abnormal karyon,alteredelectronica density,disappeard cellular membrane,cytoplasmic degradation.Futhermore,gap between cells became larger,cellular conjunction disappeared,synaptic vesicles weredisrupted,and accumulation of the neurosecretion was decreased.However,changesmentioned above were not so significant in the P.xylostella larvae parasitized by D.semiclausum.
6 Individual development of D.semiclausum:oogenesis with associated polydnavirus,external morphology and development of immature stages
The egg of D.semiclausum is hymenopteriform,lacking appendages.The larvae appearto be hypermetamorphic with four instars.The pupal period has two distinct phases:(a)phase I:appears larval-like with remarkable groups of urate cells;(b) phase II:exarate pupalappearance with a pigmentation sequence of eyes,ocelli,ma ndibles,thorax,abdomen,andantennae.
D.semiclausum female was found to carry a kind of PDVs (DsIV) as its majorparasitoid-associated factors beside venom for the first time.Protein content,most of whichwas ranged from 35 kDa to 220 kDa.in each venom reservior was 1.016μg.DslVs weretypical ichneumonid polydnaviruses with the following characteristics:( 1 ) the virus particles were assembled in the nuclei of calyx cells where they acquire an initial (inner) membrane,then migrate through the cytoplasm and budd out into the lumen of the ovary,and at thistime they acquire a second envelope (outer membrane);(2) the particle,which had adiameter of 130-150 nm with a length of 300-400 nm,has a genome comprising severalDNA segments.
Ovary of female D.semiclausum had 12 orarioles,and belonged to the polytrophic type.Ultrastructural observations demonstrated that several cells existed in a gametangium,andone of the central cells of the cluster differentiated into an oocyte while the remainingbecame nurse cells.Germ cells continued to develop in the vitellarium until mature;howeverthe nurse cells were in gradual degradation simultaneously.The calyx region was emptywhen the adult was just emerged;and after 2-3 days,mature eggs began to enter the calyxregion and surrounded by calyx fluid but without DsIVs attached.
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