蛋白激酶Prodeath-S/TK和细胞周期调控蛋白CKS1在棉铃虫生长发育中的功能研究
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摘要
研究背景、存在的科学问题及研究意义
昆虫生长发育主要受胰岛素(insulin)、蜕皮激索(20E)和保幼激素(JH)调控。Insulin对细胞的增殖和生长起着重要的调控作用,而20E和JH主要调控蜕皮与变态,三种激素间存在相互作用,共同调控生长发育及蜕皮变态。当JH存在的时候,高浓度的20E能够启始幼虫期蜕皮的发生,当JH滴度下降或消失的时候,高浓度的20E则启动昆虫变态,而insulin则通过促进幼虫生长,使前胸腺合成高浓度的蜕皮激素参与生长和变态调控。
变态期幼虫组织凋亡,成虫组织形成,其中,幼虫中肠的解体与成虫中肠的重建涉及细胞程序性死亡和细胞增殖,是目前研究的热点。蛋白激酶能通过促进底物的磷酸化来参与多种重要的生理过程,多种蛋白激酶在促进细胞程序性死亡的过程中发挥着重要的功能,CKS1作为细胞周期的重要调控蛋白,能够与CDK及Cyclin相互作用,共同调控细胞的增殖。但促细胞死亡激酶与CKS1在昆虫的生长发育及变态过程中的功能尚不清楚,Insulin、20E和JH如何调控这些基因表达也不清楚。研究CKS1和促细胞死亡蛋白激酶在昆虫生长发育及变态过程中的功能及激素调控,可以进一步阐释激素调控昆虫生长发育及变态的分子机理,为害虫防治提供理论依据和靶标基因。
研究结果与结论
本文以鳞翅目夜蛾科的棉铃虫及其表皮细胞系为材料,通过激素调控、RNA干扰、过表达及基因突变等多种方法,深入研究了CKS1和Prodeath S/TK在棉铃虫生长发育、变态及组织解体与重建过程中的功能及作用机理,获得如下结果和结论：
1. Prodeath S/TK参与变态期旧组织的程序性细胞死亡
Prodeath S/TK在蜕皮和变态期表达水平明显增加,能够被20E诱导上调表达。虫体干扰Prodeath S/TK能够抑制20E信号通路中与细胞凋亡相关的基因表达,导致旧组织解体受到抑制,严重影响昆虫变态的发生。免疫细胞化学实验结果显示,Prodeath S/TK主要是在细胞质中发挥功能,20E诱导不能使其发生亚细胞定位变化。在锦铃虫表皮细胞中过表达Prodeath S/TK能够促使细胞程序性死亡,缺失突变结果显示,S TKc结构域是Prodeath S/TK的功能域,只有当S_TKc结构域定位在细胞质中时,才能够发挥其促细胞程序性死亡的功能,而其N-末端和其C-末端决定了其亚细胞定位。蜕皮激素促进Prodeath S/TK表达,在细胞质中参与20E信号通路来行使其促细胞程序性死亡的功能。
2. CKS1通过参与insulin信号通路调控昆虫的生长发育
CKS1主要在棉铃虫幼虫生长阶段高表达,在变态期表达水平显著降低。虫体干扰CKS1能够影响insulin信号通路中与生长发育相关的基因表达,延迟生长和变态,形成体型较小的蛹Insulin和20E能够分别上调CKS1的表达水平但两种激素叠加则抑制CKS1表达。在棉铃虫表皮细胞系中过表达的CKS1主要分布在细胞核中,能够促进细胞的增殖,基因突变结果显示CKS结构域行使主要功能,其中的第45位氨基酸N45为其活性位点。虫体干扰CKS1抑制了中肠细胞增殖,进而影响生长所需的营养物质的吸收,使虫体不能达到起始变态所需的临界体重,从而阻碍了幼虫的正常生长和变态的发生。CKS1通过参与insulin信号通路促进生长阶段虫体的细胞增殖,达到临界体重产生高浓度的蜕皮激素,蜕皮激素与insulin(?)叠加终止了CKS1表达和细胞增殖,进入变态。创新点及意义
首次鉴定到一种新的蛋白激酶Prodeath S/TK,阐明了该激酶在昆虫变态期通过参与20E信号通路调控旧组织凋亡的分子机理,为20E调控变态期组织解体提供了新的实验证据和理论支持。通过对CKS1(?)的功能研究,揭示了昆早由生长转向变态的分子机理,为insulin参与调控昆虫生长与变态提供了实验依据,阐明了insulin与20E共同作用调控昆虫生长发育的机理。为昆虫发育提供了新的理论知识,为害虫防治提供的新的靶标。
Background, scientific questions and significance of researches
Insect development is regulated by insulin,20E and JH. Insulin primarily regulates cell proliferation and growth, while20E and JH coordinately regulate molting and metamorphosis. During the development process of holometabolous insect, when JH exists, high titer of20E can initiate the larval molting. When JH is absent, high titer of20E will initiate the metamorphosis. Insulin mainly promotes larval growth, and therefore the prothoracic gland (PG) growth, which produces high level of20E to initiate metamorphosis. The cross talk of three hormones orchestrates insect growth, molting and metamorphosis.
Tissue remodeling occurs during insect metamorphosis, that is the larval tissues degradation by programmed cell death (PCD), and the adult tissues formation by cell proliferation. Insect tissue remodeling presents good model to study the hormonal regulation on PCD and cell proliferation. Protein kinase participate in various important physiological processes via promote substrates phosphoralation, a set of protein kinase were reported to promote programmed cell death, CKS1as a key factor in regulating cell cycle, can interact with CDKs and Cyclins, co-regulate cell proliferation. While the functions of protein kinase and CKS1in insect development and metamorphosis are still obscure, and in which way insulin,20E and JH regulate the expression of these genes are still unclear. Deeply study the functions and hormone regulation of CKS1and Prodeath S/TK can help to understand the molecular mechanisms of insect growth and metamorphosis, provide more valuable information for further study of insect physiology and pest management.
Results and conclusion
The Lepidoptera, cotton bollworm (Helicoverpa armigera) and HaEpi (Helicoverpa epidermis cell line) were used to study the functions amd molecular mechanisms of CKS1and Prodeath S/TK in insect development and tissues remolding by various molecular methods. The results are as follows:
1. Prodeath S/TK takes part in programmed cell death (PCD) during metamorphosis
Prodeath S/TK is highly expressed at the molting and metamorphosis stages both in transcription level and in translation level It can be upregulation by20E. Knockdown of it by RNAi can suppress the expression of genes related to PCD in20E signal pathway, block the old tissues degradation and metamorphosis. Immunocytochemistry showed that Prodeath S/TK is mainly localized in cytoplasm to play its function.20E can not shift its subcellular localization. Overexpression analysis suggests CKS1can promote programmed cell death. Deletion mutations indicate S_TKc is its function domain. The N-and C-terminal determine its subcellular localization. The prodeath S/TK takes part in20E signal pathway in cytoplasm to promote cell death by20E regulation.
2. CKS1takes part in insulin signal pathway to promote cell proliferation
CKS1is highly expressed at the growth stage of insect development and is decreased at metamorphosis stage. Knockdown of CKS1in vivo by RNAi can suppress the expression of genes related to development in insulin signal pathway, block insect development and form small pupae. Both insulin and20E can upregulate the expression level of CKS1, respectively, while high titer of20E can block insulin-induced expression of CKS1. Overexpression of CKS1in HaEpi cell line can promote cell proliferation, with the subcellular localization mainly in the nucleus, and N45as its activity site. Knockdown of CKS1in vivo can block cell proliferation in midgut, affect insect growth, repress larvae reach to critical body weight, therefore block the initiation of metamorphosis. During larval growth stage, CKS1takes part in insulin signal pathway to promote cell proliferation. When larvae reach to critical body weight, high titer of20E overlaps insulin to block the expression of CKS1, then stops cell proliferation and initiate metamorphosis.
Innovations and significations
This work identified a novel protein kinase for the first time and named it as Prodeath S/TK. The prodeath S/TK participates in20E signal pathway to regulate old tissues degradation via PCD at metamorphosis stage. This work also demonstrates the function CKS1in insect development, which determines larval growth or metamorphosis by hormone regulation. These data can help to deeply understand the interaction of insulin and20E, thereby to understand the insect development and pest management.
昆虫生长发育主要受胰岛素(insulin)、蜕皮激索(20E)和保幼激素(JH)调控。Insulin对细胞的增殖和生长起着重要的调控作用,而20E和JH主要调控蜕皮与变态,三种激素间存在相互作用,共同调控生长发育及蜕皮变态。当JH存在的时候,高浓度的20E能够启始幼虫期蜕皮的发生,当JH滴度下降或消失的时候,高浓度的20E则启动昆虫变态,而insulin则通过促进幼虫生长,使前胸腺合成高浓度的蜕皮激素参与生长和变态调控。
变态期幼虫组织凋亡,成虫组织形成,其中,幼虫中肠的解体与成虫中肠的重建涉及细胞程序性死亡和细胞增殖,是目前研究的热点。蛋白激酶能通过促进底物的磷酸化来参与多种重要的生理过程,多种蛋白激酶在促进细胞程序性死亡的过程中发挥着重要的功能,CKS1作为细胞周期的重要调控蛋白,能够与CDK及Cyclin相互作用,共同调控细胞的增殖。但促细胞死亡激酶与CKS1在昆虫的生长发育及变态过程中的功能尚不清楚,Insulin、20E和JH如何调控这些基因表达也不清楚。研究CKS1和促细胞死亡蛋白激酶在昆虫生长发育及变态过程中的功能及激素调控,可以进一步阐释激素调控昆虫生长发育及变态的分子机理,为害虫防治提供理论依据和靶标基因。
研究结果与结论
本文以鳞翅目夜蛾科的棉铃虫及其表皮细胞系为材料,通过激素调控、RNA干扰、过表达及基因突变等多种方法,深入研究了CKS1和Prodeath S/TK在棉铃虫生长发育、变态及组织解体与重建过程中的功能及作用机理,获得如下结果和结论：
1. Prodeath S/TK参与变态期旧组织的程序性细胞死亡
Prodeath S/TK在蜕皮和变态期表达水平明显增加,能够被20E诱导上调表达。虫体干扰Prodeath S/TK能够抑制20E信号通路中与细胞凋亡相关的基因表达,导致旧组织解体受到抑制,严重影响昆虫变态的发生。免疫细胞化学实验结果显示,Prodeath S/TK主要是在细胞质中发挥功能,20E诱导不能使其发生亚细胞定位变化。在锦铃虫表皮细胞中过表达Prodeath S/TK能够促使细胞程序性死亡,缺失突变结果显示,S TKc结构域是Prodeath S/TK的功能域,只有当S_TKc结构域定位在细胞质中时,才能够发挥其促细胞程序性死亡的功能,而其N-末端和其C-末端决定了其亚细胞定位。蜕皮激素促进Prodeath S/TK表达,在细胞质中参与20E信号通路来行使其促细胞程序性死亡的功能。
2. CKS1通过参与insulin信号通路调控昆虫的生长发育
CKS1主要在棉铃虫幼虫生长阶段高表达,在变态期表达水平显著降低。虫体干扰CKS1能够影响insulin信号通路中与生长发育相关的基因表达,延迟生长和变态,形成体型较小的蛹Insulin和20E能够分别上调CKS1的表达水平但两种激素叠加则抑制CKS1表达。在棉铃虫表皮细胞系中过表达的CKS1主要分布在细胞核中,能够促进细胞的增殖,基因突变结果显示CKS结构域行使主要功能,其中的第45位氨基酸N45为其活性位点。虫体干扰CKS1抑制了中肠细胞增殖,进而影响生长所需的营养物质的吸收,使虫体不能达到起始变态所需的临界体重,从而阻碍了幼虫的正常生长和变态的发生。CKS1通过参与insulin信号通路促进生长阶段虫体的细胞增殖,达到临界体重产生高浓度的蜕皮激素,蜕皮激素与insulin(?)叠加终止了CKS1表达和细胞增殖,进入变态。创新点及意义
首次鉴定到一种新的蛋白激酶Prodeath S/TK,阐明了该激酶在昆虫变态期通过参与20E信号通路调控旧组织凋亡的分子机理,为20E调控变态期组织解体提供了新的实验证据和理论支持。通过对CKS1(?)的功能研究,揭示了昆早由生长转向变态的分子机理,为insulin参与调控昆虫生长与变态提供了实验依据,阐明了insulin与20E共同作用调控昆虫生长发育的机理。为昆虫发育提供了新的理论知识,为害虫防治提供的新的靶标。
Background, scientific questions and significance of researches
Insect development is regulated by insulin,20E and JH. Insulin primarily regulates cell proliferation and growth, while20E and JH coordinately regulate molting and metamorphosis. During the development process of holometabolous insect, when JH exists, high titer of20E can initiate the larval molting. When JH is absent, high titer of20E will initiate the metamorphosis. Insulin mainly promotes larval growth, and therefore the prothoracic gland (PG) growth, which produces high level of20E to initiate metamorphosis. The cross talk of three hormones orchestrates insect growth, molting and metamorphosis.
Tissue remodeling occurs during insect metamorphosis, that is the larval tissues degradation by programmed cell death (PCD), and the adult tissues formation by cell proliferation. Insect tissue remodeling presents good model to study the hormonal regulation on PCD and cell proliferation. Protein kinase participate in various important physiological processes via promote substrates phosphoralation, a set of protein kinase were reported to promote programmed cell death, CKS1as a key factor in regulating cell cycle, can interact with CDKs and Cyclins, co-regulate cell proliferation. While the functions of protein kinase and CKS1in insect development and metamorphosis are still obscure, and in which way insulin,20E and JH regulate the expression of these genes are still unclear. Deeply study the functions and hormone regulation of CKS1and Prodeath S/TK can help to understand the molecular mechanisms of insect growth and metamorphosis, provide more valuable information for further study of insect physiology and pest management.
Results and conclusion
The Lepidoptera, cotton bollworm (Helicoverpa armigera) and HaEpi (Helicoverpa epidermis cell line) were used to study the functions amd molecular mechanisms of CKS1and Prodeath S/TK in insect development and tissues remolding by various molecular methods. The results are as follows:
1. Prodeath S/TK takes part in programmed cell death (PCD) during metamorphosis
Prodeath S/TK is highly expressed at the molting and metamorphosis stages both in transcription level and in translation level It can be upregulation by20E. Knockdown of it by RNAi can suppress the expression of genes related to PCD in20E signal pathway, block the old tissues degradation and metamorphosis. Immunocytochemistry showed that Prodeath S/TK is mainly localized in cytoplasm to play its function.20E can not shift its subcellular localization. Overexpression analysis suggests CKS1can promote programmed cell death. Deletion mutations indicate S_TKc is its function domain. The N-and C-terminal determine its subcellular localization. The prodeath S/TK takes part in20E signal pathway in cytoplasm to promote cell death by20E regulation.
2. CKS1takes part in insulin signal pathway to promote cell proliferation
CKS1is highly expressed at the growth stage of insect development and is decreased at metamorphosis stage. Knockdown of CKS1in vivo by RNAi can suppress the expression of genes related to development in insulin signal pathway, block insect development and form small pupae. Both insulin and20E can upregulate the expression level of CKS1, respectively, while high titer of20E can block insulin-induced expression of CKS1. Overexpression of CKS1in HaEpi cell line can promote cell proliferation, with the subcellular localization mainly in the nucleus, and N45as its activity site. Knockdown of CKS1in vivo can block cell proliferation in midgut, affect insect growth, repress larvae reach to critical body weight, therefore block the initiation of metamorphosis. During larval growth stage, CKS1takes part in insulin signal pathway to promote cell proliferation. When larvae reach to critical body weight, high titer of20E overlaps insulin to block the expression of CKS1, then stops cell proliferation and initiate metamorphosis.
Innovations and significations
This work identified a novel protein kinase for the first time and named it as Prodeath S/TK. The prodeath S/TK participates in20E signal pathway to regulate old tissues degradation via PCD at metamorphosis stage. This work also demonstrates the function CKS1in insect development, which determines larval growth or metamorphosis by hormone regulation. These data can help to deeply understand the interaction of insulin and20E, thereby to understand the insect development and pest management.
引文
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