日本柄瘤蚜茧蜂的寄生对黑豆蚜的发育和生化代谢影响的研究
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摘要
黑豆蚜[Aphis craccivora (Koch)]属同翅目、蚜科，别名有花生蚜、苜蓿蚜、菜豆蚜、槐蚜等，是危害严重的世界性害虫，寄主植物广泛。除取食对植物造成的损害外，黑豆蚜还传播植物病毒病，导致严重的经济损失。对有害昆虫进行生物防治是控制虫害的重要手段。日本柄瘤蚜茧蜂[Lysiphlebus japonicus (Ashmead)]是黑豆蚜重要的寄生性天敌，属膜翅目，蚜茧蜂科，是营内寄生的单寄生蜂，对黑豆蚜有较好的控制作用。前人对日本柄瘤蚜茧蜂仅作过一些野外调查和简单的生物学研究，目前，对日本柄瘤蚜茧蜂的发育及成功寄生的条件和机制仍缺乏基本的信息。
本文就日本柄瘤蚜茧蜂的寄生影响因子、龄期划分及对寄主黑豆蚜的发育和生化代谢调控进行了研究，结果报告如下。
1. 本部分以黑豆蚜为寄主，研究了寄生蜂蜂龄、接蜂时间、寄生蜂密度、寄主密度、寄主龄期和翅型等因子对日本柄瘤蚜茧蜂寄生的影响。实验室研究表明，在25±1℃温度条件下，1天龄的日本柄瘤蚜茧蜂雌蜂对寄主的寄生率最高，其次为2-4天龄雌蜂，5天龄雌蜂寄生率明显降低。接蜂时间对该蜂的寄生显示明显影响，但同时又受寄主密度的影响。在接蜂时间相对短的情况下，寄生率随着接蜂时间的增加而增加。在蜂蚜比为1:20时，持续接蜂超过24小时，寄生率不再明显增长；在蜂蚜比为1:40的条件下，寄生率在持续接蜂超过48小时后才无明显增加。寄生蜂密度亦影响日本柄瘤蚜茧蜂的寄生，在蜂密度为1、5、10、15头雌蜂/200头蚜虫时，寄生率随着雌蜂密度的增加而逐渐升高，但当蜂密度增加至20、30头雌蜂/200头蚜虫时，寄生率不升反降。寄主密度对日本柄瘤蚜茧蜂的寄生有明显影响，在寄主密度为50-100头蚜虫/5头雌蜂时，寄生率最高，达90%以上，但过寄生率也高，随着寄主密度的升高，蚜虫的寄生率
逐步下降，同时过寄生率也相应逐步降低。日本柄瘤蚜茧蜂对寄主龄期显示明显的选择性，龄期较小的蚜虫寄生率较高，2龄若蚜的寄生率最高，1龄若蚜次之，而4龄若蚜的寄生率最低。在有翅与无翅型3、4龄若蚜寄主中，日本柄瘤蚜茧蜂较喜寄生无翅型若蚜，无翅型若蚜的寄生率均高于同龄有翅型若蚜。
2.为了详细了解日本柄瘤蚜茧蜂的幼虫发育，研究日本柄瘤蚜茧蜂与黑豆蚜之间发育的相互关系，我们利用扫描电镜技术区分了日本柄瘤蚜茧蜂的四个幼虫龄期。对日本柄瘤蚜茧蜂的雌蜂剖腹卵与产出卵进行了首次对比描述。这是关于日本柄瘤蚜茧蜂幼期形态的首次报道。日本柄瘤蚜茧蜂有四个幼虫龄期。1龄幼虫有发达的骨化大颚，其主要在种内的物理竞争中发挥作用。同时，幼虫还具有背部刺突和尾突。2龄幼虫大颚显著退化，背部密布小刺簇，尾突短而钝。发达的大颚在幼虫进入3龄后重新出现，此时的幼虫体壁光滑，尾部退化呈小突起。4龄幼虫大颚较3龄退化，头部器官明显分化。文中用扫描电镜照片描述了各龄期的区分特点。
3. 本部分研究了日本柄瘤蚜茧蜂对黑豆蚜龄期的选择和黑豆蚜龄期对日本柄瘤蚜茧蜂发育的影响。试验结果表明，在混合虫态寄主中，日本柄瘤蚜茧蜂通常选择黑豆蚜的较小龄期若蚜寄生，其中2龄若蚜被寄生的相对机率最高，为26.4%，其次是1龄若蚜，为20.6%，无翅成蚜与3、4龄有翅若蚜和成蚜的相对寄生机率较低。在25℃下，日本柄瘤蚜茧蜂的寄生延缓了黑豆蚜若蚜的发育，其中1龄寄生若蚜的发育在一、二、三龄被显著延滞，有翅3龄若蚜被寄生后其发育在三、四龄明显延长，但无翅和有翅4龄寄生若蚜的发育均不受影响；寄生后的各龄若蚜的成蚜寿命明显缩短，其中，1龄寄生的若蚜不能发育至成蚜，其他较早龄期寄生的若蚜生育能力则被显著抑制。在25℃下，日本柄瘤蚜茧蜂在不同龄期黑豆蚜体内发育历期有所差异，寄生2龄黑豆蚜的日本柄瘤蚜茧蜂个体发育最快，为194.10小时，在1龄寄主体内的蚜茧蜂个体发育最慢，需215.80小时。
4. 日本柄瘤蚜茧蜂的寄生影响了寄主黑豆蚜体内的生化代谢。试验选用2龄若蚜进行。寄生使黑豆蚜的游离氨基酸总浓度升高，寄生1天后，正常组的游离氨基酸总浓度为17.721nmol/L，寄生组为23.153nmol/L；寄生4天后，正常组和寄生组分别为58.703和69.659nmol/L。苏氨酸、谷氨酸和酪氨酸是黑豆蚜血淋巴中主要的氨基酸，寄生后的黑豆蚜血淋巴中苏氨酸含量升高，谷氨酸和酪氨酸含量下降。与正常蚜虫相比，寄生3天后，黑豆蚜血淋巴的蛋白质浓度下降；体蛋白质浓度升高；寄生还使寄主血淋巴的蛋白质组成发生了变化，在SDS-聚丙烯酰胺凝胶电泳图谱中出现了两个新的蛋白质，其分子量分别约为54和41kD。在寄生后4天，黑豆蚜的血淋巴海藻糖浓度明显降低。寄生蚜虫的体总糖原和体总脂含量在寄生后2天显著升高，但在寄生后3、4天明显低于未寄生蚜虫。
5. 从黑豆蚜体内经DE52 Cellulose、Sephadex G-150凝胶过滤以及FPLC Hitrap Q Sepharose分离出一种蛋白质。该蛋白质经SDS-PAGE测定在非还原和还原条件下分子量均为60kD左右，等电聚焦测得其等电点约为5.0, 为糖蛋白，不是脂蛋白。若蚜期间该蛋白质在蚜虫体内积累，蜕皮至成蚜时含量明显减少，为蚜虫提供发育所需营养，在成蚜期间仍以低浓度存在。根据其分子量和氨基酸组成与含量变化动态应
Aphis craccivora Koch (Hemiptera: Aphididae), black bean aphid or peanut aphid, is a worldwide serious pest with numerous hosts. Besides feeding damage, A. craccivora spreads plant viruses, and results in heavily economic loss. Biological control is an important measure to control pests. Lysiphlebus japonicus Ashmead (Hymenoptera: Aphidiidae), a solitary endoparasitoid, is a useful parasitic natural enemy of A. craccivora. Although some field investigations and biological features have been done, the basic information of development and the mechanism of successful parasitism still lacking in L. japonicus.
This paper studied the effects of some factors on parasitism, the determination of L. japonicus instars, the regulations on host development and metabolism. The results are as follows:
1. In the laboratory condition, parasitoid stage, exposure period, female parasitoid density, host density, host instar and morph of host wings were found to affect the parasitization of L. japonicus on A. craccivora. The parasitization rate dereased with the increase of parasitoid age. The highest parasitization rate was obtained by parasitoids of 1d age. Exposure period affected the parasitization of L. japonicus. When the parasite:host ratio was 1:20, the parasitization rates did not
increase distinctively if the exposure period was more than 24hr.. While the ratio was 1:40, the parasitization rates kept on increasing significantly until the exposure period was more than 48hr.. The parasitoid density could affect the parasitization of L. japonicus too. When the parasitoid density was 1、5、10、15 female parasitoids/200 aphids, the parasitization rate increased readily with parasitoid density. However, when the parasitoid density was 20、30 female parasitoids/5 aphids, the parasitization decreased. Host density had significant effect on the parasitism of L. japonicus. The highest parasitization rate was more than 90%, which was gained at 50-100 aphids/5 female parasitoids. Both parasitization rate and superparasitism rate decreased with the increase of host density. The host instar could influence the parasitization rate. L. japonicus preferred to oviposit on blank bean aphid of younger instar and the parasitization rate for L2 was the highest, the second highest was for L1, but that for L4 was the lowest. Between apterous and alate aphid larvae, L. japonicus preferred for apterous larvae. All parasitization rates of apterous larvae were higher than those of alate host of the same instar.
2. To make clear the larval development of L. japonicus and study the developmental interactions between L. japonicus and A. craccivora farther, we described and illustrated with scanning electron micrographs the chief diagnostic criteria of 4 instars of L. japonicus each. This is the first report on larval morph of L. japonicus. The 1st instar is mandibulate, caudate and has sclerotized spines. Supernumerary newly hatched larvae are eliminated by powerful mandibles of the prior hatched larva. The 2nd instar has degenerative mandibles and covered with small sclerotized bristles on abdominal segments; the cauda is short and blunt. The 3rd instar is mandibulate; its cuticle is smooth and the caudal segment retrogresses to a short salient. The 4th instar is strong and hymenopteriform; the mandibles are
smaller than those of 3rd instar, and various sensory organs are well developed, especially for the mouthpart.
3. In the laboratory conditions, host-instar selection of Lysiphlebus japonicus Ashmead and its development in Aphis craccivora Koch of different instars were studied. When served mixed instars aphids, The parasitoid showed a preference for small aphids. The relative parasitization rate of L2 aphids was the highest (26.4%), followed by L1 aphids, which was 20.6%, the others were lower. The devolopment of aphids was delayed after parasitized. At 25℃, the developmental period of parasitized L1 aphids was significantly prolonged at the 1st、2nd and the 3rd instar; and which of alate parasitized L3 aphids was eviden
本文就日本柄瘤蚜茧蜂的寄生影响因子、龄期划分及对寄主黑豆蚜的发育和生化代谢调控进行了研究，结果报告如下。
1. 本部分以黑豆蚜为寄主，研究了寄生蜂蜂龄、接蜂时间、寄生蜂密度、寄主密度、寄主龄期和翅型等因子对日本柄瘤蚜茧蜂寄生的影响。实验室研究表明，在25±1℃温度条件下，1天龄的日本柄瘤蚜茧蜂雌蜂对寄主的寄生率最高，其次为2-4天龄雌蜂，5天龄雌蜂寄生率明显降低。接蜂时间对该蜂的寄生显示明显影响，但同时又受寄主密度的影响。在接蜂时间相对短的情况下，寄生率随着接蜂时间的增加而增加。在蜂蚜比为1:20时，持续接蜂超过24小时，寄生率不再明显增长；在蜂蚜比为1:40的条件下，寄生率在持续接蜂超过48小时后才无明显增加。寄生蜂密度亦影响日本柄瘤蚜茧蜂的寄生，在蜂密度为1、5、10、15头雌蜂/200头蚜虫时，寄生率随着雌蜂密度的增加而逐渐升高，但当蜂密度增加至20、30头雌蜂/200头蚜虫时，寄生率不升反降。寄主密度对日本柄瘤蚜茧蜂的寄生有明显影响，在寄主密度为50-100头蚜虫/5头雌蜂时，寄生率最高，达90%以上，但过寄生率也高，随着寄主密度的升高，蚜虫的寄生率
逐步下降，同时过寄生率也相应逐步降低。日本柄瘤蚜茧蜂对寄主龄期显示明显的选择性，龄期较小的蚜虫寄生率较高，2龄若蚜的寄生率最高，1龄若蚜次之，而4龄若蚜的寄生率最低。在有翅与无翅型3、4龄若蚜寄主中，日本柄瘤蚜茧蜂较喜寄生无翅型若蚜，无翅型若蚜的寄生率均高于同龄有翅型若蚜。
2.为了详细了解日本柄瘤蚜茧蜂的幼虫发育，研究日本柄瘤蚜茧蜂与黑豆蚜之间发育的相互关系，我们利用扫描电镜技术区分了日本柄瘤蚜茧蜂的四个幼虫龄期。对日本柄瘤蚜茧蜂的雌蜂剖腹卵与产出卵进行了首次对比描述。这是关于日本柄瘤蚜茧蜂幼期形态的首次报道。日本柄瘤蚜茧蜂有四个幼虫龄期。1龄幼虫有发达的骨化大颚，其主要在种内的物理竞争中发挥作用。同时，幼虫还具有背部刺突和尾突。2龄幼虫大颚显著退化，背部密布小刺簇，尾突短而钝。发达的大颚在幼虫进入3龄后重新出现，此时的幼虫体壁光滑，尾部退化呈小突起。4龄幼虫大颚较3龄退化，头部器官明显分化。文中用扫描电镜照片描述了各龄期的区分特点。
3. 本部分研究了日本柄瘤蚜茧蜂对黑豆蚜龄期的选择和黑豆蚜龄期对日本柄瘤蚜茧蜂发育的影响。试验结果表明，在混合虫态寄主中，日本柄瘤蚜茧蜂通常选择黑豆蚜的较小龄期若蚜寄生，其中2龄若蚜被寄生的相对机率最高，为26.4%，其次是1龄若蚜，为20.6%，无翅成蚜与3、4龄有翅若蚜和成蚜的相对寄生机率较低。在25℃下，日本柄瘤蚜茧蜂的寄生延缓了黑豆蚜若蚜的发育，其中1龄寄生若蚜的发育在一、二、三龄被显著延滞，有翅3龄若蚜被寄生后其发育在三、四龄明显延长，但无翅和有翅4龄寄生若蚜的发育均不受影响；寄生后的各龄若蚜的成蚜寿命明显缩短，其中，1龄寄生的若蚜不能发育至成蚜，其他较早龄期寄生的若蚜生育能力则被显著抑制。在25℃下，日本柄瘤蚜茧蜂在不同龄期黑豆蚜体内发育历期有所差异，寄生2龄黑豆蚜的日本柄瘤蚜茧蜂个体发育最快，为194.10小时，在1龄寄主体内的蚜茧蜂个体发育最慢，需215.80小时。
4. 日本柄瘤蚜茧蜂的寄生影响了寄主黑豆蚜体内的生化代谢。试验选用2龄若蚜进行。寄生使黑豆蚜的游离氨基酸总浓度升高，寄生1天后，正常组的游离氨基酸总浓度为17.721nmol/L，寄生组为23.153nmol/L；寄生4天后，正常组和寄生组分别为58.703和69.659nmol/L。苏氨酸、谷氨酸和酪氨酸是黑豆蚜血淋巴中主要的氨基酸，寄生后的黑豆蚜血淋巴中苏氨酸含量升高，谷氨酸和酪氨酸含量下降。与正常蚜虫相比，寄生3天后，黑豆蚜血淋巴的蛋白质浓度下降；体蛋白质浓度升高；寄生还使寄主血淋巴的蛋白质组成发生了变化，在SDS-聚丙烯酰胺凝胶电泳图谱中出现了两个新的蛋白质，其分子量分别约为54和41kD。在寄生后4天，黑豆蚜的血淋巴海藻糖浓度明显降低。寄生蚜虫的体总糖原和体总脂含量在寄生后2天显著升高，但在寄生后3、4天明显低于未寄生蚜虫。
5. 从黑豆蚜体内经DE52 Cellulose、Sephadex G-150凝胶过滤以及FPLC Hitrap Q Sepharose分离出一种蛋白质。该蛋白质经SDS-PAGE测定在非还原和还原条件下分子量均为60kD左右，等电聚焦测得其等电点约为5.0, 为糖蛋白，不是脂蛋白。若蚜期间该蛋白质在蚜虫体内积累，蜕皮至成蚜时含量明显减少，为蚜虫提供发育所需营养，在成蚜期间仍以低浓度存在。根据其分子量和氨基酸组成与含量变化动态应
Aphis craccivora Koch (Hemiptera: Aphididae), black bean aphid or peanut aphid, is a worldwide serious pest with numerous hosts. Besides feeding damage, A. craccivora spreads plant viruses, and results in heavily economic loss. Biological control is an important measure to control pests. Lysiphlebus japonicus Ashmead (Hymenoptera: Aphidiidae), a solitary endoparasitoid, is a useful parasitic natural enemy of A. craccivora. Although some field investigations and biological features have been done, the basic information of development and the mechanism of successful parasitism still lacking in L. japonicus.
This paper studied the effects of some factors on parasitism, the determination of L. japonicus instars, the regulations on host development and metabolism. The results are as follows:
1. In the laboratory condition, parasitoid stage, exposure period, female parasitoid density, host density, host instar and morph of host wings were found to affect the parasitization of L. japonicus on A. craccivora. The parasitization rate dereased with the increase of parasitoid age. The highest parasitization rate was obtained by parasitoids of 1d age. Exposure period affected the parasitization of L. japonicus. When the parasite:host ratio was 1:20, the parasitization rates did not
increase distinctively if the exposure period was more than 24hr.. While the ratio was 1:40, the parasitization rates kept on increasing significantly until the exposure period was more than 48hr.. The parasitoid density could affect the parasitization of L. japonicus too. When the parasitoid density was 1、5、10、15 female parasitoids/200 aphids, the parasitization rate increased readily with parasitoid density. However, when the parasitoid density was 20、30 female parasitoids/5 aphids, the parasitization decreased. Host density had significant effect on the parasitism of L. japonicus. The highest parasitization rate was more than 90%, which was gained at 50-100 aphids/5 female parasitoids. Both parasitization rate and superparasitism rate decreased with the increase of host density. The host instar could influence the parasitization rate. L. japonicus preferred to oviposit on blank bean aphid of younger instar and the parasitization rate for L2 was the highest, the second highest was for L1, but that for L4 was the lowest. Between apterous and alate aphid larvae, L. japonicus preferred for apterous larvae. All parasitization rates of apterous larvae were higher than those of alate host of the same instar.
2. To make clear the larval development of L. japonicus and study the developmental interactions between L. japonicus and A. craccivora farther, we described and illustrated with scanning electron micrographs the chief diagnostic criteria of 4 instars of L. japonicus each. This is the first report on larval morph of L. japonicus. The 1st instar is mandibulate, caudate and has sclerotized spines. Supernumerary newly hatched larvae are eliminated by powerful mandibles of the prior hatched larva. The 2nd instar has degenerative mandibles and covered with small sclerotized bristles on abdominal segments; the cauda is short and blunt. The 3rd instar is mandibulate; its cuticle is smooth and the caudal segment retrogresses to a short salient. The 4th instar is strong and hymenopteriform; the mandibles are
smaller than those of 3rd instar, and various sensory organs are well developed, especially for the mouthpart.
3. In the laboratory conditions, host-instar selection of Lysiphlebus japonicus Ashmead and its development in Aphis craccivora Koch of different instars were studied. When served mixed instars aphids, The parasitoid showed a preference for small aphids. The relative parasitization rate of L2 aphids was the highest (26.4%), followed by L1 aphids, which was 20.6%, the others were lower. The devolopment of aphids was delayed after parasitized. At 25℃, the developmental period of parasitized L1 aphids was significantly prolonged at the 1st、2nd and the 3rd instar; and which of alate parasitized L3 aphids was eviden
引文
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