两种小GTP酶Rab4b，Rab32及蛋白激酶C delta（PKC δ）在棉铃虫变态过程中的功能研究
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摘要
研究背景
昆虫是地球上种类最多,数量最大的生物种群,与人类有着密不可分的关系。昆虫的发育是由激素调控的复杂的生命过程,包括昆虫的蜕皮与变态。在全变态昆虫中,蜕皮过程包括幼虫之间的蜕皮,幼虫到蛹的蜕皮以及蛹到成虫的蜕皮。幼虫向蛹的转变被称为变态。昆虫的变态是受蜕皮激素(20E)和保幼激素(JH)共同调控的。在幼虫期,20E引发蜕皮,JH负责维持个体的幼虫状态；而在幼虫的末龄期,JH滴度逐渐降低甚至消失,20E则起始变态过程。目前对于20E及JH协调调控变态的机制已经有了比较深入的研究。但最近的研究表明,除了20E与JH两种激素外,胰岛素(insulin)在昆虫的变态中起着不可忽视的作用。研究发现,insulin与20E存在着相互作用,二者共同的调控昆虫的生长及变态决定。但二者相互作用的分子机制尚不清楚,有哪些分子参与了两个通路的相互作用,还有待进一步研究。
昆虫的变态过程中伴随着组织的重建,例如,中肠和脂肪体。在变态时期,幼虫的组织要发生解体,成虫的组织重新形成,这一过程也是由20E与JH共同调控的。20E诱导程序性细胞死亡使幼虫组织降解,而JH则拮抗20E的功能。然而,组织的重建过程是一个复杂的生理过程,除了激素的调控外,还依赖于其他的活性分子。但目前对于参与组织重建效应分子的研究还不深入。小GTP酶Rab蛋白在哺乳动物中广泛参与了各种生理活动,包括膜泡运输,细胞骨架活性及信号转导等生理过程,但在昆虫发育过程中对于该类蛋白的研究还很有限。此外,蛋白激酶C作为一种丝氨酸/苏氨酸蛋白激酶可以通过促进底物蛋白磷酸化调节多种生理过程,如细胞生长与增殖,细胞凋亡等。有研究发现蛋白激酶C参与昆虫20E信号转导过程,但哪种蛋白激酶C参与20E的信号转导还有待研究。
存在的科学问题
本实验室通过对棉铃虫变态期转录组进行测序,鉴定到两种小GTP酶Rab4b,Rab32及一种蛋白激酶C delta (PKCδ)基因。拟解决的科学问题为：Rab4b, Rab32是否参与由激素调控的变态过程?在功能上有何差异?PKCδ是否参与20E调节的变态及其伴随的组织重建过程?
研究意义
本研究通过鉴定三种参与变态过程的关键分子,阐明了不同分子的具体功能,从分子水平上揭示激素调控变态的分子机制,为进一步研究昆虫发育提供依据。此外,本研究通过鉴定参与组织重建的效应分子,丰富了我们对于组织重建机制的认识和理解,为农业害虫的防治提供了理论指导。
研究方案及取得的成果
本文以鳞翅目昆虫,棉铃虫,为实验材料,通过体内,体外的干扰实验及激素调控实验来探索两种Rab蛋白Rab4b,Rab32及蛋白激酶C delta (PKCδ)在昆虫变态及组织重建中的功能。所得实验结果如下：
1.Rab4b参与insulin与20E信号通路的基因转录进而调控糖原水平及变态发生
本研究在棉铃虫鉴定到一个小GTP酶Rab4b参与了20E与胰岛素调控的基因转录。该基因在表皮、中肠、脂肪体及血淋巴中均有表达。虫体RNA干扰导致幼虫在化蛹前死亡,异常化蛹及小蛹的形成。生化分析结果显示,干扰Rab4b后,糖原的储存水平下降；体内及体外干扰实验表明,20E通路中的Br和HR3以及糖原合成酶GS的转录水平受到抑制,而FOXO的转录水平上调。免疫细胞化学及免疫印迹的结果表明,insulin通过Rab4b抑制FOXO的核定位,过表达Rab4b进行亚细胞定位检测,发现insulin没有影响Rab4b定位,而20E刺激后Rab4b向细胞膜迁移。但在mRNA水平上,20E不影响Rab4b的表达,而胰岛素则诱导Rab4b的上调。暗示20E与胰岛素信号通路通过不同的方式来调控Rab4b,最终调控变态的发生。
2.Rab32参与棉铃虫变态期中肠的重建
我们从棉铃虫中肠中鉴定到Rab32基因。半定量RT-PCR分析Rab32表达模式发现该基因在预蛹期的中肠mRNA水平最高。20E诱导Rab32上调,而JH类似物(JHA)不影响其转录。在表皮细胞系中干扰20E核受体EcR-B1后,20E对Rab32的上调作用消除。免疫细胞化学结果显示,该蛋白主要位于细胞质中,而20E使Rab32在细胞系中聚集颗粒状结构并增加Rab32在细胞质中的蛋白表达。免疫组织化学结果显示,在变态期的中肠中,Rab32定位与成虫中肠上皮细胞的内部边缘位置,暗示其可能参与中肠形成中营养的吸收及信号转导的作用。在虫体内干扰Rab32后,幼虫的生长及变态没有受到影响,但成虫中肠的形态发生异常。这些结果说明Rab32参与了棉铃虫变态期的中肠重建,Rab32可能在20E信号途径的调控下参与成虫中肠的形成。
3.蛋白激酶C delta(PKCδ)参与棉铃虫变态期组织的解体
本研究在棉铃虫中发现了一个新型的蛋白激酶C,PKCδ,参与了变态期中肠和脂肪体的凋亡。PKCδ在棉铃虫的表皮、中肠、脂肪体中均有表达,且在变态期转录水平较高。体内注射激素实验结果显示,20E和JHⅢ都可以上调PKCδ的转录,。在细胞系上干扰EcR-B1后,20E不能上调PKCδ,而干扰JH的候选受体Met后,JHⅢ对PKCδ的诱导作用没有变化,说明20E通过EcR0B1调控PKCδ,而JHⅢ可能通过其他的方式,而不是Met来调控PKCδ转录。在虫体中干扰PKCδ,中肠和脂肪体无法进行解体和重建；凋亡相关基因Caspase8的表达被抑制,而凋亡抑制因子Survivin转录上调。在表皮细胞中过表达PKCδ的功能域,可以诱导细胞凋亡,并增强了FOXO的核定位,说明PKCδ可能通过调节FOXO的转录活性诱导凋亡。在表皮细胞中干扰PKCδ,抑制了20E及JHⅢ对USPl,Calponin等关键基因的上调作用,说明PKCδ可能参与了20E与JH的信号转导途径的相互作用。
所取得成果的创新点与意义
本文主要通过体内及体外的RNA干扰技术,鉴定到一个小GTP酶Rab4b通过调控20E与胰岛素通路基因转录及糖原合成进而参与变态。深入揭示了20E通路与胰岛素通路相互作用的机制。同时,我们鉴定到另一种小GTP酶Rab32参与了中肠重建中成虫中肠的形成,丰富了小GTP酶Rab蛋白在昆虫中的功能研究。此外,发现蛋白激酶C delta(PKCδ)参与了20E诱导的组织凋亡及20E与JH对下游基因的调控,本研究为深入揭示昆虫变态的激素调控及组织重建的分子机制提供了依据。
Background
Insects are the largest biological species with a maximum population on earth, which have a close relationship with humans. Insect development is a complicated physiological process regulated by hormones, including molting and metamorphosis. In holometabolous insects, insect molting contains molt from larvae-larvae, larvae-pupa and pupa-adult. The larva to pupae transition is called metamorphosis. This process is controlled by two kinds of hormones:20E and JH. At larval stage,20E initiate molting, and JH maintains larval growth. At the final larval stage,20E provokes metamorphosis when JH level decreases. Previous studies have uncovered the mechanism of20E and JH regulated metamorphic transition. Whereas, recent finding indicates that insulin play a crucial role in larvae-pupae transform. Insulin could cooperate with20E signal and regulating insect development. However, the molecular mechanism that insulin interacts with20E during insect metamorphosis is not fully understood.
During insect metamorphosis, tissues undergo remodeling under the control of20E and JH. In this process, the larval tissues, such as midgut and fat body undergo degradation by20E induced programmed cell death (PCD), and then the new tissues form by imaginal cell proliferation, whereas, JH antagonizes20E effects. The foregoing investigations have revealed the hormone regulations on tissue remodeling. Nevertheless, molecules involved in this process need to be further identified. In mammalian, the Small GTPases are widely found to take part in multiple physiological processes, such as visecle trafficking, cytoskeleton activity and signal transduction. Howerve, the function of Rab proteins are not well studied in insect. The protein kinase C is also reported to play important roles in cell growth, proliferation and apoptosis. Recently studies indicate the protein particiapates in20E signaling.
Pending Problems
What are the functions of Rab4b and Rab32in hormones regulated metamorphosis? And how is the difference between Rab4b and Rab32? Whether PKCδ in invovoled in20E pathway and tissue remodeling during larval-pupal transition.
Significance of the Research
The rapid development of molecular technologies brings us a convenient platform for further study the insect development. Insect undergoes molting and metamorphosis during its life. The larval-pupal transition is very crucial process for insect mature and reproduction, which is mainly regulated by20E and JH. However, more and more researches find that insulin signaling play vital role in metamorphosis by interacting with20E pathway. Thus, questions remained. The mechanism of the crosstalk between20E and insulin pathway is not well understood. Therefore, it is very meaningful to have a better conmand of the interaction of the two pathways. Moreover, tissue remodeling is an important physiological process during metamorphosis. Charactiaziton of the effetors involved in the process is with great importance and providing new thought for pest-cpntrol
Methods and Acquired Results
In the current study, we characterized a small GTPase Rab4b, participates in metamorphosis by regulating20E and insulin signals. It provided a target molecule for further understand the crosstalk between20E and insulin pathway. Moreover, we found another Rab protein, Rab32, is involved in imaginal midgut formation. Besides, our study demonstrated PKC delta (PKCδ) takes part in20E induced tissue degradation through cell death. Our investigations provide a theoretical basis for further reveal the molecular mechanism of metamorphosis and tissue reconstruction.
1. Rab4b participates in metamorphosis by regulating glycogen level and gene transcription insulin and20E signaling.
In the present study, we found that a small GTPase Rab4b from a lepidopteran insect participates in signal transduction in the two pathways. RT-PCR results showed that Rab4b was present consistently in various tissues during larval development. Knocked down of Rab4b caused53%larvae to die before pupation,24%failed to pass the metamorphic transition and became malformed pupae, and part of thelarvae transformed to smaller pupae with lower body weight. These results indicate that Rab4b is critical to larval survival and metamorphosis. Glycogen levels declined in the larval body after Rab4b silencing. Knockdown of Rab4b suppressed the gene transcription in the20E (Br and HR3) and insulin pathways (GS), whereas, FOXO increased after Rab4b interference, RNAi in HaEpi cells confirmed the results in vivo. Immunocytochemistry and western bolt showed that insulin kept FOXO located in the cytoplasm after1h treatment, whereas FOXO could not be arrested in the cytoplasm after insulin induction after knockdown of Rab4b. Overexpression experiment revealed that insulin could not alter the cytoplasm location of Rab4b. However, the20E induced Rab4b moving toward the cell membrane. Hormone regulation results indicate that20E did not effect the transcription of Rab4b, but insulin enhanced the transcription of Rab4b in3to12h after injection. Besides, the Rab4b transcription depended on the glucose level.
2. Rab32and the remodeling of the imaginal midgut in Helicoverpa armigera
We got the cDNA sequence of Rab32, RT-PCR results show that Rab32is up-regulated in epidermis and midgut during metamorphosis. Its expression could be up-regulated by20E, but not by JHA. RNAi in HaEpi cells demonstrated Rab32could be regulated by20E signal transduction pathway through EcRB1as a receptor of20E. Immunocytochemistry showed that Rab32mostly existed in the cytoplasm in normal condition,20E induced Rab32accumulation in the cytosolic granular astructures. Western blot showed20E enhanced Rab32in cytoplasm. Immunohistochemistry results revealed that the Rab32signal was detected in the cytoplasm of midgut cells in both the5th feeding larvae and metamorphic committed larvae. In the5th feeding larval midgut, the intensive signal was detected around the basement of the midgut and the apical region of the epithelium. In the metamorphic committed midgut at the6th120h larval stage, the most intensive signal was detected at the apical region of the epithelium cells of the imaginal midgut, suggesting Rab32may relate to imaginal midgut formation by nutrient absorption or signal transduction. Knockdown of Rab32 expression disrupts the imaginal midgut morphogenesis during metamorphosis. These data suggest that Rab32takes part in imaginal midgut formation.
3. Protein Kinase C delta participates in tissue degradation during metamorphosis in Helicoverpa armigera
Here we find a novel type Protein Kinase C δ (PKCδ) takes part in the apoptosis in midgut and the fat body in a lepidopteran insect Helicoverpa armigera. PKCδ was highly expressed in several tissues during metamorphic stage, and could be stimulated by20E and Juvenile hormone (JHⅢ). Knockdown of EcR-B1in HaEpi cells blocked20E induced PKCδ transcription, but interference of JH candidate receptor, Met, did not effect the JHIII induced PKCδ transcription, suggesting20E regulates PKCδ through EcR-B1, whereas JH effects PKCδ via other way except Met. Immunohistochemistry and HE staining indicated that the fat body and the midgut did not experience degradation in the PKCδ knockdown larvae. RT-PCR analysis demonstrated the apoptosis related gene Caspase8transcription was blocked after PKCδ knockdown, whereas, the inhibitor of apoptosis, Survivin, was enhanced in the RNAi larvae. The data suggest PKCδ participate in the hydrolyzing of the midgut and the fat body during metamorphosis. Overexpress functional domain of PKCδ in HaEpi cells induced apoptosis, and the indicator of apoptosis Caspase8was upregulated, meanwhile the IAP, Survivin was inhibited, which further evidenced the active role of PKCδ in HaEpi cells apoptosis. In PKCδ overexpression cells, FOXO displayed much stronger signal in the nuclei comparing with that in the normal cells. RFP was also used as control to confirm the effect of PKCδ on subcellular location of FOXO. The data suggest the overexpressed PKCδ functional domain enhance the nuclear location of FOXO. RNAi of PKCδ in HaEpi cells surpressed the20E and JHIII induced expression of USP1and Calponin, Which suggest that PKCδ involved in the20E and JH signal transduction.
昆虫是地球上种类最多,数量最大的生物种群,与人类有着密不可分的关系。昆虫的发育是由激素调控的复杂的生命过程,包括昆虫的蜕皮与变态。在全变态昆虫中,蜕皮过程包括幼虫之间的蜕皮,幼虫到蛹的蜕皮以及蛹到成虫的蜕皮。幼虫向蛹的转变被称为变态。昆虫的变态是受蜕皮激素(20E)和保幼激素(JH)共同调控的。在幼虫期,20E引发蜕皮,JH负责维持个体的幼虫状态；而在幼虫的末龄期,JH滴度逐渐降低甚至消失,20E则起始变态过程。目前对于20E及JH协调调控变态的机制已经有了比较深入的研究。但最近的研究表明,除了20E与JH两种激素外,胰岛素(insulin)在昆虫的变态中起着不可忽视的作用。研究发现,insulin与20E存在着相互作用,二者共同的调控昆虫的生长及变态决定。但二者相互作用的分子机制尚不清楚,有哪些分子参与了两个通路的相互作用,还有待进一步研究。
昆虫的变态过程中伴随着组织的重建,例如,中肠和脂肪体。在变态时期,幼虫的组织要发生解体,成虫的组织重新形成,这一过程也是由20E与JH共同调控的。20E诱导程序性细胞死亡使幼虫组织降解,而JH则拮抗20E的功能。然而,组织的重建过程是一个复杂的生理过程,除了激素的调控外,还依赖于其他的活性分子。但目前对于参与组织重建效应分子的研究还不深入。小GTP酶Rab蛋白在哺乳动物中广泛参与了各种生理活动,包括膜泡运输,细胞骨架活性及信号转导等生理过程,但在昆虫发育过程中对于该类蛋白的研究还很有限。此外,蛋白激酶C作为一种丝氨酸/苏氨酸蛋白激酶可以通过促进底物蛋白磷酸化调节多种生理过程,如细胞生长与增殖,细胞凋亡等。有研究发现蛋白激酶C参与昆虫20E信号转导过程,但哪种蛋白激酶C参与20E的信号转导还有待研究。
存在的科学问题
本实验室通过对棉铃虫变态期转录组进行测序,鉴定到两种小GTP酶Rab4b,Rab32及一种蛋白激酶C delta (PKCδ)基因。拟解决的科学问题为：Rab4b, Rab32是否参与由激素调控的变态过程?在功能上有何差异?PKCδ是否参与20E调节的变态及其伴随的组织重建过程?
研究意义
本研究通过鉴定三种参与变态过程的关键分子,阐明了不同分子的具体功能,从分子水平上揭示激素调控变态的分子机制,为进一步研究昆虫发育提供依据。此外,本研究通过鉴定参与组织重建的效应分子,丰富了我们对于组织重建机制的认识和理解,为农业害虫的防治提供了理论指导。
研究方案及取得的成果
本文以鳞翅目昆虫,棉铃虫,为实验材料,通过体内,体外的干扰实验及激素调控实验来探索两种Rab蛋白Rab4b,Rab32及蛋白激酶C delta (PKCδ)在昆虫变态及组织重建中的功能。所得实验结果如下：
1.Rab4b参与insulin与20E信号通路的基因转录进而调控糖原水平及变态发生
本研究在棉铃虫鉴定到一个小GTP酶Rab4b参与了20E与胰岛素调控的基因转录。该基因在表皮、中肠、脂肪体及血淋巴中均有表达。虫体RNA干扰导致幼虫在化蛹前死亡,异常化蛹及小蛹的形成。生化分析结果显示,干扰Rab4b后,糖原的储存水平下降；体内及体外干扰实验表明,20E通路中的Br和HR3以及糖原合成酶GS的转录水平受到抑制,而FOXO的转录水平上调。免疫细胞化学及免疫印迹的结果表明,insulin通过Rab4b抑制FOXO的核定位,过表达Rab4b进行亚细胞定位检测,发现insulin没有影响Rab4b定位,而20E刺激后Rab4b向细胞膜迁移。但在mRNA水平上,20E不影响Rab4b的表达,而胰岛素则诱导Rab4b的上调。暗示20E与胰岛素信号通路通过不同的方式来调控Rab4b,最终调控变态的发生。
2.Rab32参与棉铃虫变态期中肠的重建
我们从棉铃虫中肠中鉴定到Rab32基因。半定量RT-PCR分析Rab32表达模式发现该基因在预蛹期的中肠mRNA水平最高。20E诱导Rab32上调,而JH类似物(JHA)不影响其转录。在表皮细胞系中干扰20E核受体EcR-B1后,20E对Rab32的上调作用消除。免疫细胞化学结果显示,该蛋白主要位于细胞质中,而20E使Rab32在细胞系中聚集颗粒状结构并增加Rab32在细胞质中的蛋白表达。免疫组织化学结果显示,在变态期的中肠中,Rab32定位与成虫中肠上皮细胞的内部边缘位置,暗示其可能参与中肠形成中营养的吸收及信号转导的作用。在虫体内干扰Rab32后,幼虫的生长及变态没有受到影响,但成虫中肠的形态发生异常。这些结果说明Rab32参与了棉铃虫变态期的中肠重建,Rab32可能在20E信号途径的调控下参与成虫中肠的形成。
3.蛋白激酶C delta(PKCδ)参与棉铃虫变态期组织的解体
本研究在棉铃虫中发现了一个新型的蛋白激酶C,PKCδ,参与了变态期中肠和脂肪体的凋亡。PKCδ在棉铃虫的表皮、中肠、脂肪体中均有表达,且在变态期转录水平较高。体内注射激素实验结果显示,20E和JHⅢ都可以上调PKCδ的转录,。在细胞系上干扰EcR-B1后,20E不能上调PKCδ,而干扰JH的候选受体Met后,JHⅢ对PKCδ的诱导作用没有变化,说明20E通过EcR0B1调控PKCδ,而JHⅢ可能通过其他的方式,而不是Met来调控PKCδ转录。在虫体中干扰PKCδ,中肠和脂肪体无法进行解体和重建；凋亡相关基因Caspase8的表达被抑制,而凋亡抑制因子Survivin转录上调。在表皮细胞中过表达PKCδ的功能域,可以诱导细胞凋亡,并增强了FOXO的核定位,说明PKCδ可能通过调节FOXO的转录活性诱导凋亡。在表皮细胞中干扰PKCδ,抑制了20E及JHⅢ对USPl,Calponin等关键基因的上调作用,说明PKCδ可能参与了20E与JH的信号转导途径的相互作用。
所取得成果的创新点与意义
本文主要通过体内及体外的RNA干扰技术,鉴定到一个小GTP酶Rab4b通过调控20E与胰岛素通路基因转录及糖原合成进而参与变态。深入揭示了20E通路与胰岛素通路相互作用的机制。同时,我们鉴定到另一种小GTP酶Rab32参与了中肠重建中成虫中肠的形成,丰富了小GTP酶Rab蛋白在昆虫中的功能研究。此外,发现蛋白激酶C delta(PKCδ)参与了20E诱导的组织凋亡及20E与JH对下游基因的调控,本研究为深入揭示昆虫变态的激素调控及组织重建的分子机制提供了依据。
Background
Insects are the largest biological species with a maximum population on earth, which have a close relationship with humans. Insect development is a complicated physiological process regulated by hormones, including molting and metamorphosis. In holometabolous insects, insect molting contains molt from larvae-larvae, larvae-pupa and pupa-adult. The larva to pupae transition is called metamorphosis. This process is controlled by two kinds of hormones:20E and JH. At larval stage,20E initiate molting, and JH maintains larval growth. At the final larval stage,20E provokes metamorphosis when JH level decreases. Previous studies have uncovered the mechanism of20E and JH regulated metamorphic transition. Whereas, recent finding indicates that insulin play a crucial role in larvae-pupae transform. Insulin could cooperate with20E signal and regulating insect development. However, the molecular mechanism that insulin interacts with20E during insect metamorphosis is not fully understood.
During insect metamorphosis, tissues undergo remodeling under the control of20E and JH. In this process, the larval tissues, such as midgut and fat body undergo degradation by20E induced programmed cell death (PCD), and then the new tissues form by imaginal cell proliferation, whereas, JH antagonizes20E effects. The foregoing investigations have revealed the hormone regulations on tissue remodeling. Nevertheless, molecules involved in this process need to be further identified. In mammalian, the Small GTPases are widely found to take part in multiple physiological processes, such as visecle trafficking, cytoskeleton activity and signal transduction. Howerve, the function of Rab proteins are not well studied in insect. The protein kinase C is also reported to play important roles in cell growth, proliferation and apoptosis. Recently studies indicate the protein particiapates in20E signaling.
Pending Problems
What are the functions of Rab4b and Rab32in hormones regulated metamorphosis? And how is the difference between Rab4b and Rab32? Whether PKCδ in invovoled in20E pathway and tissue remodeling during larval-pupal transition.
Significance of the Research
The rapid development of molecular technologies brings us a convenient platform for further study the insect development. Insect undergoes molting and metamorphosis during its life. The larval-pupal transition is very crucial process for insect mature and reproduction, which is mainly regulated by20E and JH. However, more and more researches find that insulin signaling play vital role in metamorphosis by interacting with20E pathway. Thus, questions remained. The mechanism of the crosstalk between20E and insulin pathway is not well understood. Therefore, it is very meaningful to have a better conmand of the interaction of the two pathways. Moreover, tissue remodeling is an important physiological process during metamorphosis. Charactiaziton of the effetors involved in the process is with great importance and providing new thought for pest-cpntrol
Methods and Acquired Results
In the current study, we characterized a small GTPase Rab4b, participates in metamorphosis by regulating20E and insulin signals. It provided a target molecule for further understand the crosstalk between20E and insulin pathway. Moreover, we found another Rab protein, Rab32, is involved in imaginal midgut formation. Besides, our study demonstrated PKC delta (PKCδ) takes part in20E induced tissue degradation through cell death. Our investigations provide a theoretical basis for further reveal the molecular mechanism of metamorphosis and tissue reconstruction.
1. Rab4b participates in metamorphosis by regulating glycogen level and gene transcription insulin and20E signaling.
In the present study, we found that a small GTPase Rab4b from a lepidopteran insect participates in signal transduction in the two pathways. RT-PCR results showed that Rab4b was present consistently in various tissues during larval development. Knocked down of Rab4b caused53%larvae to die before pupation,24%failed to pass the metamorphic transition and became malformed pupae, and part of thelarvae transformed to smaller pupae with lower body weight. These results indicate that Rab4b is critical to larval survival and metamorphosis. Glycogen levels declined in the larval body after Rab4b silencing. Knockdown of Rab4b suppressed the gene transcription in the20E (Br and HR3) and insulin pathways (GS), whereas, FOXO increased after Rab4b interference, RNAi in HaEpi cells confirmed the results in vivo. Immunocytochemistry and western bolt showed that insulin kept FOXO located in the cytoplasm after1h treatment, whereas FOXO could not be arrested in the cytoplasm after insulin induction after knockdown of Rab4b. Overexpression experiment revealed that insulin could not alter the cytoplasm location of Rab4b. However, the20E induced Rab4b moving toward the cell membrane. Hormone regulation results indicate that20E did not effect the transcription of Rab4b, but insulin enhanced the transcription of Rab4b in3to12h after injection. Besides, the Rab4b transcription depended on the glucose level.
2. Rab32and the remodeling of the imaginal midgut in Helicoverpa armigera
We got the cDNA sequence of Rab32, RT-PCR results show that Rab32is up-regulated in epidermis and midgut during metamorphosis. Its expression could be up-regulated by20E, but not by JHA. RNAi in HaEpi cells demonstrated Rab32could be regulated by20E signal transduction pathway through EcRB1as a receptor of20E. Immunocytochemistry showed that Rab32mostly existed in the cytoplasm in normal condition,20E induced Rab32accumulation in the cytosolic granular astructures. Western blot showed20E enhanced Rab32in cytoplasm. Immunohistochemistry results revealed that the Rab32signal was detected in the cytoplasm of midgut cells in both the5th feeding larvae and metamorphic committed larvae. In the5th feeding larval midgut, the intensive signal was detected around the basement of the midgut and the apical region of the epithelium. In the metamorphic committed midgut at the6th120h larval stage, the most intensive signal was detected at the apical region of the epithelium cells of the imaginal midgut, suggesting Rab32may relate to imaginal midgut formation by nutrient absorption or signal transduction. Knockdown of Rab32 expression disrupts the imaginal midgut morphogenesis during metamorphosis. These data suggest that Rab32takes part in imaginal midgut formation.
3. Protein Kinase C delta participates in tissue degradation during metamorphosis in Helicoverpa armigera
Here we find a novel type Protein Kinase C δ (PKCδ) takes part in the apoptosis in midgut and the fat body in a lepidopteran insect Helicoverpa armigera. PKCδ was highly expressed in several tissues during metamorphic stage, and could be stimulated by20E and Juvenile hormone (JHⅢ). Knockdown of EcR-B1in HaEpi cells blocked20E induced PKCδ transcription, but interference of JH candidate receptor, Met, did not effect the JHIII induced PKCδ transcription, suggesting20E regulates PKCδ through EcR-B1, whereas JH effects PKCδ via other way except Met. Immunohistochemistry and HE staining indicated that the fat body and the midgut did not experience degradation in the PKCδ knockdown larvae. RT-PCR analysis demonstrated the apoptosis related gene Caspase8transcription was blocked after PKCδ knockdown, whereas, the inhibitor of apoptosis, Survivin, was enhanced in the RNAi larvae. The data suggest PKCδ participate in the hydrolyzing of the midgut and the fat body during metamorphosis. Overexpress functional domain of PKCδ in HaEpi cells induced apoptosis, and the indicator of apoptosis Caspase8was upregulated, meanwhile the IAP, Survivin was inhibited, which further evidenced the active role of PKCδ in HaEpi cells apoptosis. In PKCδ overexpression cells, FOXO displayed much stronger signal in the nuclei comparing with that in the normal cells. RFP was also used as control to confirm the effect of PKCδ on subcellular location of FOXO. The data suggest the overexpressed PKCδ functional domain enhance the nuclear location of FOXO. RNAi of PKCδ in HaEpi cells surpressed the20E and JHIII induced expression of USP1and Calponin, Which suggest that PKCδ involved in the20E and JH signal transduction.
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