菜蛾绒茧蜂滞育诱导过程中的母代效应
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摘要
菜蛾绒茧蜂Cotesia vestalis (Haliday)是小菜蛾Plutella xylostella (Linnaeus)幼虫期的主要内寄生蜂,该蜂经历不同光周期会显著影响子代的滞育。对菜蛾绒茧蜂滞育诱导过程中的母代效应进行深入研究,对于提高该蜂的滞育诱导技术,进一步利用滞育特性,解决实际应用过程中面临的长期储藏等问题有着重要的现实意义。鉴于此,本文对饲养于25±1℃8L：16D、12L：12D和16L：8D三种光周期下(G0)以及子代(G1)饲养于13℃8L：16D下的菜蛾绒茧蜂物质代谢进行研究,结果如下：
一、光周期对菜蛾绒茧蜂形态发生、个体大小及发育的影响
菜蛾绒茧蜂的生活史分为卵、幼虫、蛹和成虫4个虫态,在幼虫的末期存在一个形态上明显可区别于幼虫早期的发育阶段(预蛹)。光周期对各发育阶段的发育历期无显著影响,在25±1℃,三种光周期下,菜蛾绒茧蜂胚胎发育期、1龄、2龄和3龄幼虫期、预蛹期以及蛹期均分别为36h、36h、84h、12h、24h和108h,但光周期显著影响虫体大小。胚胎体积随发育时间逐渐增大,胚胎发育在16h至32h期间,其体积在三种不同光照下存在不规则差异,但到孵化时三种光周期下的胚胎体积无显著差异。1龄幼虫及其尾针长、宽8h短光照下的均大于16h长光照下的,但大多数发育时间差异并不显著。2龄幼虫长、宽8h短光照下的大于16h长光照下的,在96h,108h,120h和144h发育时间点,尾囊长、宽均显著大于长光照下的。3龄幼虫及其尾囊长、宽8h短光照下的与16h长光照下的无显著差异。G,代滞育预蛹的体积显著大于Go代非滞育预蛹的体积。180h的预蛹及茧的体积均以饲养于8h短光照下的显著大于16h长光照下的,尾囊长、宽则均以16h长光照下的最大。192h的预蛹体积在8h和16h光周期下无显著的差异。羽化前一天,8h短光照下的蛹长、前胸背板宽以及腹宽均大于16h长光照下的。羽化后,雌成虫长、前胸背板宽和翅展及其腹部体积均随饲养光周期的延长而减小。三种光周期下饲养获得的成虫所怀的卵(产出前)和产出后的子代卵体积均以8h短光照下饲养的最大。
二、光周期对菜蛾绒茧蜂重量及含水量的影响
G1代滞育预蛹的鲜重,干重均显著重于G0代的非滞育预蛹。Go代预蛹和雌成虫的鲜重和干重均随饲养光照的延长而下降。
含水量G1代滞育预蛹与G0代的非滞育预蛹无显著差异。G0代预蛹和雌成虫含水量均随饲养光周期的延长而升高。结合水含量G1代滞育预蛹显著高于Go代的非滞育预蛹。8h和12h光照下的预蛹结合水含量显著高于16h光照下的。雌成虫结合水含量随饲养光照的延长而下降。
三、菜蛾绒茧蜂滞育诱导过程中过氧化氢代谢的母代效应
过氧化氢(hydrogen peroxide, H2O2)含量在G0代预蛹期和成虫期均随饲养光周期的延长而升高,8h短光照饲养的Go代预蛹与G1代滞育预蛹无显著差异,但G1代卵在产出前和产出后均以饲养于8h短光照下的最高。过氧化物歧化酶(superoxide dismutase, SOD)和过氧化物酶(peroxidase, POD)活性在G0代预蛹期,成虫期及G1代卵在产出前和产出后均随饲养光周期的延长而上升,而过氧化氢酶(catalase, CAT)活性则随饲养光周期的延长显著降低。与G1代滞育预蛹相比,8h短光照下的G0代预蛹SOD活性显著增高,而CAT活性则显著降低,POD活性无显著差异。在三种光周期下G1代卵无论在产出前还是在产出后,三种保护酶活性的变化趋势均与G0代的预蛹期和成虫期相似。但是,POD和CAT的活性在G1代卵产出后显著高于卵产出前。
四、菜蛾绒茧蜂滞育诱导过程中脂类代谢的母代效应
总脂滴度G1代滞育预蛹与三种光周期下的G0代非滞育预蛹之间均无显著差异。G0代预蛹总脂滴度在三种光周期之间无显著差异。G0代雌成虫总脂滴度8h短光照下显著高于16h光照。G1代卵总脂滴度在产出前随光周期的延长而降低,在产出后,8h和16h光照之间无显著差异,但均显著高于12h光照。
甘油三酯含量G1代滞育预蛹显著高于8h和16h光照下的G0代非滞育预蛹,但显著低于12h光照下的非滞育预蛹。G0代预蛹、雌成虫以及G1代卵产出前甘油三酯含量8h短光照下的显著高于16h光照下的。G1代卵产出后在三种光周期间无显著差异。
游离脂肪酸含量G1代滞育预蛹显著低于三种光周期下的G。代非滞育预蛹。G0代预蛹、G,代产出前和产出后的卵游离脂肪酸含量8h短光照下的显著低于16h长光照下的。但G0代雌成虫游离脂肪酸含量在三种光周期间无显著差异。
五、菜蛾绒茧蜂滞育诱导过程中糖类代谢的母代效应
总糖含量在8h光照下G0代幼虫、预蛹和蛹中均显著高于16h光照下的。三种光周期下成虫期总糖含量无显著差异。G1代卵、幼虫和预蛹内总糖含量随G0代饲养光照的延长而降低,且幼虫和预蛹在三种G0代饲养光周期间的差异比G0代进一步增大。
海藻糖含量,除G1代幼虫饲养于12h光周期下的显著高于16h光照下的外,其他所有测试阶段在三种光周期间均无显著差异。
在G0代,幼虫、预蛹、蛹和雌成虫糖原含量均随饲养光周期的延长而降低,但雄成虫在三种光周期间无显著差异；在G1代,卵、幼虫与预蛹的糖原含量也均随G0代饲养光周期的延长而降低,且预蛹糖原含量在三种G0代饲养光周期间的差异比G0代进一步增大。
六、菜蛾绒茧蜂滞育诱导过程中糖原合成酶和糖原磷酸化酶的母代效应
糖原合成酶活性除了G0代蛹和雄成虫在饲养于8h光照与16h光照间无显著差异之外,其它测试虫态均以G0代饲养于8h光照下的显著高于饲养于16h光照下的。糖原磷酸化酶活性除了G0代饲养于8h光照下的G1代卵期显著高于16h光照下的,G1代预蛹期显著低于16h光照下的,其它测试阶段在Go代饲养于8h和16h光照下间均无显著差异。各发育阶段糖原合成酶和糖原磷酸化酶的mRNA表达量高低与活性高低相一致。糖原合成酶mRNA表达量G0代饲养于8h光照下的均明显高于16h光照下的,而糖原磷酸化酶mRNA表达量在大多数发育阶段G0代饲养于8h光照下和16h光照下之间均无明显的差异。
七、菜蛾绒茧蜂滞育诱导过程中激素的母代效应
20-羟基蜕皮酮(20-hydroxyecdysone,20E)和保幼激素Ⅲ (juvenile hormone,JH Ⅲ)在所测定的发育阶段中均有表达。20E含量滞育预蛹显著低于非滞育预蛹,而JH Ⅲ含量与饲养于相同光照下的非滞育预蛹无显著差异。G0代预蛹和成虫20E含量随光周期的延长而升高。G1代卵(产出前和产出后),20E含量均以饲养于8h短光照下最低。在G0代成虫期和G1代卵产出后,JH Ⅲ含量三种光周期间均无显著差异。
简言之,菜蛾绒茧蜂母代经历的光周期显著影响虫体的代谢变化。短光照引起20E含量的下降,在20E的调控下,糖原合成酶基因表达上调,使活性上升导致虫体糖原储备的上升。雌成虫以H2O2作为光周期的传递因子,将大量的H2O2和糖原通过卵细胞质传递到子代,子代在其后续的发育过程中,对从母代继承的这种物质代谢信息和环境信息作出确认,并进一步强化对不利环境的反应,通过20E的持续调控,在糖原合成酶和磷酸化酶的作用下,使糖原积累的差异进一步扩大,最终影响子代预蛹进入滞育的比例。
Cotesia vestalis (Haliday) is one of the most important solitary larval endoparasitoid of Plutella xylostella (Linnaeus). Photoperiod influences diapause beyond the maternal generation in C. vestalis. It is of momentous current significance to study profoundly maternal effects on diapause induction in C. vestalis for the improvement of the technique of diapause induction, better utilization of the diapause characteristics, and solution of the long-term storage problem faced in the practical application process. Hence, material metabolism was studied at various stages in Go reared at25±1℃under photoperiods (light:dark) of8L:16D,12L:12D, and16L:8D, and in G1reared at13℃under L8:16D. The major results are summarized as follows:
1Photoperiodic effect on morphogenesis, size, and development in C. vestalis
The life history in C. vestalis is divided into four developmental stages, which are egg, larval, pupal and adult stages, and at the end of larval stage, there exists a developmental stage (prepupa), which obviously differs from early larvae in morphology. No significant influence of photoperiod on the developmental duration at each developmental stage was detected. The duration for embryonic development (egg stage), for the first, second and third instar larvae, prepupal and pupal stages in C. vestalis was36h,36h,84h,12h,24h, and108h, respectively, at25±1℃under three photoperiods. However, photoperiod had significant effect on body size. Embryo volume in Go increased gradually with the developmental time. The embryo volume at16-32h showed irregular difference among three photoperiods, but at hatching, it came to the same level. At almost all developmental time, body and caudal needle sizes of the first instar larvae under8h light were larger than those under16h light, but did not show statistically differences. The size of the second instar larvae under8h light was larger than that under16h light, especially the size of caudal vesicula at96h,108h,120h and144h. Body size and the size of caudal vesicula in the third instar larvae under8h of short photoperiod were no significant differences compared to those under16h light. The volumes in Gi diapausing prepupae and cocoons were significantly higher than those in Go nondiapausing prepupae and cocoons. At180h, the volumes in Go prepupae and cocoons under8h light were significantly higher than those under16h light, whereas the size of caudal vesicula under16h light was the largest. At192h, there was no significant difference in volume of prepupae between under8h and under16h light. The size of pupa at last day was larger under8h light compared to that under16h light. After emergence, size, and abdominal volume in adult females decreased as the light period was prolonged. The volumes in Gi eggs both before and after oviposition under8h light were the largest.
2Photoperiodic effect on weight, and water content in C. vestalis
Fresh and dry weights were significantly heavier in G1diapausing prepupae than those in Go nondiapausing prepupae. Fresh and dry weights decreased at both the Go nondiapausing prepupal and adult stages as the light period was extended.
No significant difference was detected in water content between Go nondiapausing prepupae and G1diapausing prepupae. Water content at both the Go nondiapausing prepupal and adult stages increased as the light period was prolonged. G1diapausing prepupae had significantly higher bound water content than Go nondiapausing prepupae.G0nondiapausing prepupae reared under8h and12h light had significantly higher bound water content than that reared under16h of long-light. Moreover, bound water content in the Go adult females decreased with increased photoperiod.
3Maternal effect on hydrogen peroxide metabolism during diapause induction in C. vestalis
The hydrogen peroxide contents in Go prepupae and adults increased as the light period was prolonged, and there was no significant difference between Go nondiapausing prepupae and G1diapausing prepupae, however, the hydrogen peroxide content in G1eggs both before and after oviposition under8h short light was the highest. Activities of superoxide dismutase and peroxidase in Go prepupae, adults, and G1eggs both before and after oviposition increased, whereas catalase activity decreased significantly as the light period was prolonged. Compared to G1diapausing prepupae, Go prepupae under8h short light had significantly higher superoxide dismutase, lower catalase, and similar peroxidase activities. Under the three photoperiods, the variation trends of protective enzyme activities in G1eggs (before and after oviposition) were similar to the trends in Go prepupae and adults. Peroxidase and catalase activities in eggs were significantly higher after oviposition than before oviposition.
4Maternal effect on lipid metabolism during diapause induction in C. vestalis
Total lipid titer between G1diapausing prepupae and Go nondiapausing prepupae under any photoperiod had no significant difference. Total lipid titer in the Go prepupae showed no significant differences among three photoperiods, either. Total lipid titer in Go adult females under8h light was significantly higher than that under16h light. Total lipid titer in G1eggs before oviposition was decreased with increased photoperiod. After oviposition, the total lipid titer in G1egg under8h light was similar to that under16h light, but both were significantly higher than that under12h light.
Triglyceride content in G1diapausing prepupae was significantly higher than that in Go nondiapausing prepupae both under8h and16h light, but lower than that under12h light. Triglyceride contents in Go prepupae, adult females and G1egg before oviposition under8h light were significantly higher than those under16h light. After oviposition, however, there was no significant difference among G1eggs under three photoperiods.
Free fatty acid content in G1diapausing prepupae was significantly lower than that in Go nondiapausing prepupae under three photoperiods. Free fatty acid contents in Go prepupae, and G1eggs (both before and after oviposition) under8h light were significantly lower than those under16h light. However, free fatty acid contents in the Go adult females among three photoperiods showed no significant differences.
5Maternal effect on carbohydrate metabolism during diapause induction in C. vestalis
Total sugar contents in Go larvae, prepupae and pupae were significantly higher under8h light than under16h light. Total sugar content in Go adults among photoperiods showed no significant difference. In G1, the total sugar contents of eggs, larvae and prepupae decreased with increased photoperiods. Furthermore, the disparity of total sugar content across the three photoperiods in G1larvae and prepupae was much bigger than that in Go larvae and prepupae.
There were no significant differences in trehalose content at all developmental stages but G1larvae, in which the content was significantly higher under12h light than that under16h light.
In Go, the glycogen contents of larvae, prepupae, pupae and adult females decreased as the light period was extended, but those of adult males did not show significant difference among the three photoperiods. In G1, the glycogen contents of eggs, larvae and prepupae also decreased with increased photoperiod. Compared to that in Go, the difference of glycogen accumulation in the prepupae of G1was expanded among three photoperiods.
6Maternal effect on glycogen synthase and phosphorylase during diapause induction in C. vestalis
Glycogen synthase activities at all tested developmental stages of both in Go and G1were significantly higher under8h light than under16h light, with the exception of those in the pupae and adult males of Go, which both under8h light and under16h light had similar glycogen synthase activities. There were no significant differences in glycogen phosphorylase activity in Go and in the larvae of G1between under8h light and under16h light. However, the eggs in G1produced by the females which were reared under8h light had significantly higher, and the prepupae in G1produced by the females which were reared under8h light had significantly lower glycogen phosphorylase activities than those produced by the females which were reared under16h light. The activities of both two enzymes showed identical to mRNA expressions of the two enzymes at all stages both in Go and G1. Glycogen synthase mRNA expressions at all stages which or whose Go were reared under8h light were obviously higher than under16h light, whereas glycogen phosphorylase mRNA expressions were similar to16h light.
7Maternal effect on hormones during diapause induction in C. vestalis
20-hydroxyecdysone (20E) and juvenile hormone Ⅲ (JH Ⅲ) could be detected at all the tested stages.20E content in diapausing prepupae was significantly lower than that in nondiapausing prepupae, whereas JH Ⅲ content did not significantly differ from that in nondiapausing prepupae under the same photoperiod. The20E content in Go prepupae and adult females increased with increased photoperiods. In G1eggs (before and after oviposition),20E contents were the lowest under8h short light. JH Ⅲ contents in both Go adult females and G1eggs (after oviposition) under three photoperiods showed no significant differences.
In brief, photoperiods experienced by Go significantly affect metabolism. The content of20E declined with decreased photoperiods, and glycogen was accumulated in C. vestalis in consequence of up-regulated gene expression and enhanced activity of glycogen synthase under the control of20E. H2O2as a factor transmitting information of experienced photoperiod, and glycogen are transferred to offspring through the cytoplasm. Moreover, information from the maternal generation is incorporated by offspring according to environmental conditions experienced by themselves. Ultimately, this information influences glycogen synthase and phosphorylase gene expressions and activities by continuous regulation of20E, thereby determining the proportion of progeny that enter diapause.
一、光周期对菜蛾绒茧蜂形态发生、个体大小及发育的影响
菜蛾绒茧蜂的生活史分为卵、幼虫、蛹和成虫4个虫态,在幼虫的末期存在一个形态上明显可区别于幼虫早期的发育阶段(预蛹)。光周期对各发育阶段的发育历期无显著影响,在25±1℃,三种光周期下,菜蛾绒茧蜂胚胎发育期、1龄、2龄和3龄幼虫期、预蛹期以及蛹期均分别为36h、36h、84h、12h、24h和108h,但光周期显著影响虫体大小。胚胎体积随发育时间逐渐增大,胚胎发育在16h至32h期间,其体积在三种不同光照下存在不规则差异,但到孵化时三种光周期下的胚胎体积无显著差异。1龄幼虫及其尾针长、宽8h短光照下的均大于16h长光照下的,但大多数发育时间差异并不显著。2龄幼虫长、宽8h短光照下的大于16h长光照下的,在96h,108h,120h和144h发育时间点,尾囊长、宽均显著大于长光照下的。3龄幼虫及其尾囊长、宽8h短光照下的与16h长光照下的无显著差异。G,代滞育预蛹的体积显著大于Go代非滞育预蛹的体积。180h的预蛹及茧的体积均以饲养于8h短光照下的显著大于16h长光照下的,尾囊长、宽则均以16h长光照下的最大。192h的预蛹体积在8h和16h光周期下无显著的差异。羽化前一天,8h短光照下的蛹长、前胸背板宽以及腹宽均大于16h长光照下的。羽化后,雌成虫长、前胸背板宽和翅展及其腹部体积均随饲养光周期的延长而减小。三种光周期下饲养获得的成虫所怀的卵(产出前)和产出后的子代卵体积均以8h短光照下饲养的最大。
二、光周期对菜蛾绒茧蜂重量及含水量的影响
G1代滞育预蛹的鲜重,干重均显著重于G0代的非滞育预蛹。Go代预蛹和雌成虫的鲜重和干重均随饲养光照的延长而下降。
含水量G1代滞育预蛹与G0代的非滞育预蛹无显著差异。G0代预蛹和雌成虫含水量均随饲养光周期的延长而升高。结合水含量G1代滞育预蛹显著高于Go代的非滞育预蛹。8h和12h光照下的预蛹结合水含量显著高于16h光照下的。雌成虫结合水含量随饲养光照的延长而下降。
三、菜蛾绒茧蜂滞育诱导过程中过氧化氢代谢的母代效应
过氧化氢(hydrogen peroxide, H2O2)含量在G0代预蛹期和成虫期均随饲养光周期的延长而升高,8h短光照饲养的Go代预蛹与G1代滞育预蛹无显著差异,但G1代卵在产出前和产出后均以饲养于8h短光照下的最高。过氧化物歧化酶(superoxide dismutase, SOD)和过氧化物酶(peroxidase, POD)活性在G0代预蛹期,成虫期及G1代卵在产出前和产出后均随饲养光周期的延长而上升,而过氧化氢酶(catalase, CAT)活性则随饲养光周期的延长显著降低。与G1代滞育预蛹相比,8h短光照下的G0代预蛹SOD活性显著增高,而CAT活性则显著降低,POD活性无显著差异。在三种光周期下G1代卵无论在产出前还是在产出后,三种保护酶活性的变化趋势均与G0代的预蛹期和成虫期相似。但是,POD和CAT的活性在G1代卵产出后显著高于卵产出前。
四、菜蛾绒茧蜂滞育诱导过程中脂类代谢的母代效应
总脂滴度G1代滞育预蛹与三种光周期下的G0代非滞育预蛹之间均无显著差异。G0代预蛹总脂滴度在三种光周期之间无显著差异。G0代雌成虫总脂滴度8h短光照下显著高于16h光照。G1代卵总脂滴度在产出前随光周期的延长而降低,在产出后,8h和16h光照之间无显著差异,但均显著高于12h光照。
甘油三酯含量G1代滞育预蛹显著高于8h和16h光照下的G0代非滞育预蛹,但显著低于12h光照下的非滞育预蛹。G0代预蛹、雌成虫以及G1代卵产出前甘油三酯含量8h短光照下的显著高于16h光照下的。G1代卵产出后在三种光周期间无显著差异。
游离脂肪酸含量G1代滞育预蛹显著低于三种光周期下的G。代非滞育预蛹。G0代预蛹、G,代产出前和产出后的卵游离脂肪酸含量8h短光照下的显著低于16h长光照下的。但G0代雌成虫游离脂肪酸含量在三种光周期间无显著差异。
五、菜蛾绒茧蜂滞育诱导过程中糖类代谢的母代效应
总糖含量在8h光照下G0代幼虫、预蛹和蛹中均显著高于16h光照下的。三种光周期下成虫期总糖含量无显著差异。G1代卵、幼虫和预蛹内总糖含量随G0代饲养光照的延长而降低,且幼虫和预蛹在三种G0代饲养光周期间的差异比G0代进一步增大。
海藻糖含量,除G1代幼虫饲养于12h光周期下的显著高于16h光照下的外,其他所有测试阶段在三种光周期间均无显著差异。
在G0代,幼虫、预蛹、蛹和雌成虫糖原含量均随饲养光周期的延长而降低,但雄成虫在三种光周期间无显著差异；在G1代,卵、幼虫与预蛹的糖原含量也均随G0代饲养光周期的延长而降低,且预蛹糖原含量在三种G0代饲养光周期间的差异比G0代进一步增大。
六、菜蛾绒茧蜂滞育诱导过程中糖原合成酶和糖原磷酸化酶的母代效应
糖原合成酶活性除了G0代蛹和雄成虫在饲养于8h光照与16h光照间无显著差异之外,其它测试虫态均以G0代饲养于8h光照下的显著高于饲养于16h光照下的。糖原磷酸化酶活性除了G0代饲养于8h光照下的G1代卵期显著高于16h光照下的,G1代预蛹期显著低于16h光照下的,其它测试阶段在Go代饲养于8h和16h光照下间均无显著差异。各发育阶段糖原合成酶和糖原磷酸化酶的mRNA表达量高低与活性高低相一致。糖原合成酶mRNA表达量G0代饲养于8h光照下的均明显高于16h光照下的,而糖原磷酸化酶mRNA表达量在大多数发育阶段G0代饲养于8h光照下和16h光照下之间均无明显的差异。
七、菜蛾绒茧蜂滞育诱导过程中激素的母代效应
20-羟基蜕皮酮(20-hydroxyecdysone,20E)和保幼激素Ⅲ (juvenile hormone,JH Ⅲ)在所测定的发育阶段中均有表达。20E含量滞育预蛹显著低于非滞育预蛹,而JH Ⅲ含量与饲养于相同光照下的非滞育预蛹无显著差异。G0代预蛹和成虫20E含量随光周期的延长而升高。G1代卵(产出前和产出后),20E含量均以饲养于8h短光照下最低。在G0代成虫期和G1代卵产出后,JH Ⅲ含量三种光周期间均无显著差异。
简言之,菜蛾绒茧蜂母代经历的光周期显著影响虫体的代谢变化。短光照引起20E含量的下降,在20E的调控下,糖原合成酶基因表达上调,使活性上升导致虫体糖原储备的上升。雌成虫以H2O2作为光周期的传递因子,将大量的H2O2和糖原通过卵细胞质传递到子代,子代在其后续的发育过程中,对从母代继承的这种物质代谢信息和环境信息作出确认,并进一步强化对不利环境的反应,通过20E的持续调控,在糖原合成酶和磷酸化酶的作用下,使糖原积累的差异进一步扩大,最终影响子代预蛹进入滞育的比例。
Cotesia vestalis (Haliday) is one of the most important solitary larval endoparasitoid of Plutella xylostella (Linnaeus). Photoperiod influences diapause beyond the maternal generation in C. vestalis. It is of momentous current significance to study profoundly maternal effects on diapause induction in C. vestalis for the improvement of the technique of diapause induction, better utilization of the diapause characteristics, and solution of the long-term storage problem faced in the practical application process. Hence, material metabolism was studied at various stages in Go reared at25±1℃under photoperiods (light:dark) of8L:16D,12L:12D, and16L:8D, and in G1reared at13℃under L8:16D. The major results are summarized as follows:
1Photoperiodic effect on morphogenesis, size, and development in C. vestalis
The life history in C. vestalis is divided into four developmental stages, which are egg, larval, pupal and adult stages, and at the end of larval stage, there exists a developmental stage (prepupa), which obviously differs from early larvae in morphology. No significant influence of photoperiod on the developmental duration at each developmental stage was detected. The duration for embryonic development (egg stage), for the first, second and third instar larvae, prepupal and pupal stages in C. vestalis was36h,36h,84h,12h,24h, and108h, respectively, at25±1℃under three photoperiods. However, photoperiod had significant effect on body size. Embryo volume in Go increased gradually with the developmental time. The embryo volume at16-32h showed irregular difference among three photoperiods, but at hatching, it came to the same level. At almost all developmental time, body and caudal needle sizes of the first instar larvae under8h light were larger than those under16h light, but did not show statistically differences. The size of the second instar larvae under8h light was larger than that under16h light, especially the size of caudal vesicula at96h,108h,120h and144h. Body size and the size of caudal vesicula in the third instar larvae under8h of short photoperiod were no significant differences compared to those under16h light. The volumes in Gi diapausing prepupae and cocoons were significantly higher than those in Go nondiapausing prepupae and cocoons. At180h, the volumes in Go prepupae and cocoons under8h light were significantly higher than those under16h light, whereas the size of caudal vesicula under16h light was the largest. At192h, there was no significant difference in volume of prepupae between under8h and under16h light. The size of pupa at last day was larger under8h light compared to that under16h light. After emergence, size, and abdominal volume in adult females decreased as the light period was prolonged. The volumes in Gi eggs both before and after oviposition under8h light were the largest.
2Photoperiodic effect on weight, and water content in C. vestalis
Fresh and dry weights were significantly heavier in G1diapausing prepupae than those in Go nondiapausing prepupae. Fresh and dry weights decreased at both the Go nondiapausing prepupal and adult stages as the light period was extended.
No significant difference was detected in water content between Go nondiapausing prepupae and G1diapausing prepupae. Water content at both the Go nondiapausing prepupal and adult stages increased as the light period was prolonged. G1diapausing prepupae had significantly higher bound water content than Go nondiapausing prepupae.G0nondiapausing prepupae reared under8h and12h light had significantly higher bound water content than that reared under16h of long-light. Moreover, bound water content in the Go adult females decreased with increased photoperiod.
3Maternal effect on hydrogen peroxide metabolism during diapause induction in C. vestalis
The hydrogen peroxide contents in Go prepupae and adults increased as the light period was prolonged, and there was no significant difference between Go nondiapausing prepupae and G1diapausing prepupae, however, the hydrogen peroxide content in G1eggs both before and after oviposition under8h short light was the highest. Activities of superoxide dismutase and peroxidase in Go prepupae, adults, and G1eggs both before and after oviposition increased, whereas catalase activity decreased significantly as the light period was prolonged. Compared to G1diapausing prepupae, Go prepupae under8h short light had significantly higher superoxide dismutase, lower catalase, and similar peroxidase activities. Under the three photoperiods, the variation trends of protective enzyme activities in G1eggs (before and after oviposition) were similar to the trends in Go prepupae and adults. Peroxidase and catalase activities in eggs were significantly higher after oviposition than before oviposition.
4Maternal effect on lipid metabolism during diapause induction in C. vestalis
Total lipid titer between G1diapausing prepupae and Go nondiapausing prepupae under any photoperiod had no significant difference. Total lipid titer in the Go prepupae showed no significant differences among three photoperiods, either. Total lipid titer in Go adult females under8h light was significantly higher than that under16h light. Total lipid titer in G1eggs before oviposition was decreased with increased photoperiod. After oviposition, the total lipid titer in G1egg under8h light was similar to that under16h light, but both were significantly higher than that under12h light.
Triglyceride content in G1diapausing prepupae was significantly higher than that in Go nondiapausing prepupae both under8h and16h light, but lower than that under12h light. Triglyceride contents in Go prepupae, adult females and G1egg before oviposition under8h light were significantly higher than those under16h light. After oviposition, however, there was no significant difference among G1eggs under three photoperiods.
Free fatty acid content in G1diapausing prepupae was significantly lower than that in Go nondiapausing prepupae under three photoperiods. Free fatty acid contents in Go prepupae, and G1eggs (both before and after oviposition) under8h light were significantly lower than those under16h light. However, free fatty acid contents in the Go adult females among three photoperiods showed no significant differences.
5Maternal effect on carbohydrate metabolism during diapause induction in C. vestalis
Total sugar contents in Go larvae, prepupae and pupae were significantly higher under8h light than under16h light. Total sugar content in Go adults among photoperiods showed no significant difference. In G1, the total sugar contents of eggs, larvae and prepupae decreased with increased photoperiods. Furthermore, the disparity of total sugar content across the three photoperiods in G1larvae and prepupae was much bigger than that in Go larvae and prepupae.
There were no significant differences in trehalose content at all developmental stages but G1larvae, in which the content was significantly higher under12h light than that under16h light.
In Go, the glycogen contents of larvae, prepupae, pupae and adult females decreased as the light period was extended, but those of adult males did not show significant difference among the three photoperiods. In G1, the glycogen contents of eggs, larvae and prepupae also decreased with increased photoperiod. Compared to that in Go, the difference of glycogen accumulation in the prepupae of G1was expanded among three photoperiods.
6Maternal effect on glycogen synthase and phosphorylase during diapause induction in C. vestalis
Glycogen synthase activities at all tested developmental stages of both in Go and G1were significantly higher under8h light than under16h light, with the exception of those in the pupae and adult males of Go, which both under8h light and under16h light had similar glycogen synthase activities. There were no significant differences in glycogen phosphorylase activity in Go and in the larvae of G1between under8h light and under16h light. However, the eggs in G1produced by the females which were reared under8h light had significantly higher, and the prepupae in G1produced by the females which were reared under8h light had significantly lower glycogen phosphorylase activities than those produced by the females which were reared under16h light. The activities of both two enzymes showed identical to mRNA expressions of the two enzymes at all stages both in Go and G1. Glycogen synthase mRNA expressions at all stages which or whose Go were reared under8h light were obviously higher than under16h light, whereas glycogen phosphorylase mRNA expressions were similar to16h light.
7Maternal effect on hormones during diapause induction in C. vestalis
20-hydroxyecdysone (20E) and juvenile hormone Ⅲ (JH Ⅲ) could be detected at all the tested stages.20E content in diapausing prepupae was significantly lower than that in nondiapausing prepupae, whereas JH Ⅲ content did not significantly differ from that in nondiapausing prepupae under the same photoperiod. The20E content in Go prepupae and adult females increased with increased photoperiods. In G1eggs (before and after oviposition),20E contents were the lowest under8h short light. JH Ⅲ contents in both Go adult females and G1eggs (after oviposition) under three photoperiods showed no significant differences.
In brief, photoperiods experienced by Go significantly affect metabolism. The content of20E declined with decreased photoperiods, and glycogen was accumulated in C. vestalis in consequence of up-regulated gene expression and enhanced activity of glycogen synthase under the control of20E. H2O2as a factor transmitting information of experienced photoperiod, and glycogen are transferred to offspring through the cytoplasm. Moreover, information from the maternal generation is incorporated by offspring according to environmental conditions experienced by themselves. Ultimately, this information influences glycogen synthase and phosphorylase gene expressions and activities by continuous regulation of20E, thereby determining the proportion of progeny that enter diapause.
引文
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