昆虫眠性调控相关基因的研究进展
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  • 英文篇名:A Review on Molting-related Genes in Insect
  • 作者:毕思敏 ; 王平阳 ; 赵巧玲
  • 英文作者:Bi Simin;Wang Pingyang;Zhao Qiaoling;Jiangsu Key Laboratory of Sericultrual Biology and Biotechnology, College of Biotechnology, Jiangsu University of Science and Technology;Key Laboratory of Genetic Improvement of Silkworm and Mulberry, Ministry of Agriculture and Rural Affairs, Sericultural Research Institute, Chinese Academy of Agricultural Sci-ences;
  • 关键词:眠性相关基因 ; 昆虫 ; 蜕皮激素 ; 保幼激素 ; 神经肽
  • 英文关键词:Molting-related genes;;Insect;;Ecdysterone;;Juvenile hormone;;Neuropeptide
  • 中文刊名:CYKE
  • 英文刊名:Science of Sericulture
  • 机构:江苏科技大学生物技术学院江苏省蚕桑生物学与生物技术重点实验室;中国农业科学院蚕业研究所农业农村部桑蚕遗传改良重点实验室;
  • 出版日期:2019-04-09 17:31
  • 出版单位:蚕业科学
  • 年:2019
  • 期:v.45
  • 基金:国家自然科学基金项目(No.31372378);; 江苏省研究生创新计划项目(No.KYCX17-1857);; 江苏科技大学研究生创新计划项目(No.YCX16B-02)
  • 语种:中文;
  • 页:CYKE201901018
  • 页数:12
  • CN:01
  • ISSN:32-1115/S
  • 分类号:130-141
摘要
入眠是昆虫幼虫生长发育过程中非常重要的一种生理现象,其入眠蜕皮过程主要包括促前胸腺激素(PTTH)介导的蜕皮激素合成、蜕皮激素介导的信号转导和最终皮层溶离及色素沉积3个阶段,是一个复杂的生理过程,受到许多因素的影响,其中蜕皮激素和保幼激素等因素的影响较为显著。基于此,本文论述了蜕皮激素信号通路相关基因、保幼激素合成与代谢相关基因、相关神经肽基因及其他相关基因对昆虫眠性的影响。通过对这些基因进行综述,可以为系统研究家蚕等昆虫入眠蜕皮过程中的生理生化反应提供理论依据,对昆虫生长发育和生殖的研究以及农林害虫防控都有十分重要的意义。
        Molting is one of the important physiological processes during the growth and development of insect larva. The molting process has three stages including the synthesis of ecdysone mediated by prothoracicotropic hormone(PTTH), the signal transduction mediated by ecdysone, and cortical dissociation and pigmentation. It is a complex physiological process and affected by many factors, among which ecdysone and juvenile hormone are the most important factors. Based on these facts, we summarized the effect of a series of genes on molting in insect, including genes involved in ecdysone signalling pathway, in juvenile hormone synthesis and metabolism pathways, and neuropeptide genes as well as other related genes. This review not only provides a theoretical basis for systematically study on physiological and biochemical process of molting in silkworm and other insects, but also plays an important role in the study of growth and reproduction of insect, as well as prevention and control of the agricultural and forestal pest.
引文
[1]KLOWDEN M J.Physiological systems in insects[M].Idaho:Acdemic Press,2013:77-79
    [2]GORMAN M J,ARAKANE Y.Tyrosine hydroxylase is required for cuticle sclerotization and pigmentation in Tribolium castaneum[J].Insect Biochem Mol Biol,2010,40(3):267-273
    [3]SUGUMARAN M.Complexities of cuticular pigmentation in insects[J].Pigment Cell Melanoma Res,2009,22(5):523-525
    [4]TALLOEN W,VAN DYCK H,LENS L.The cost of melanization:butterfly wing coloration under environmental stress[J].Evolution,2004,58(2):360-366
    [5]MESCE K A,FAHRBACH S E.Integration of endocrine signals that regulate insect ecdysis[J].Front Neuroendocrinol,2002,23(2):179-199
    [6]贺伟强,刘淑梅,时连根.家蚕的眠性及其调控[J].生物学通报,2003,38(6):20-21
    [7]YAMANAKA N,REWITZ K F,O'CONNOR M B.Ecdysone control of developmental transitions:lessons from Drosophila research[J].Annu Rev Entomol,2013,58:497-516
    [8]赵小凡.昆虫蜕皮的分子机理及应用[J].应用昆虫学报,2007,44(3):323-326
    [9]NIWA R,NAMIKI T,ITO K,et al.Non-molting glossy/shroud encodes a short-chain dehydrogenase/reductase that functions in the'Black Box'of the ecdysteroid biosynthesis pathway[J].Development,2010,137(12):1991-1999
    [10]NIWA Y S,NIWA R.Neural control of steroid hormone biosynthesis during development in the fruit fly Drosophila melanogaster[J].Genes Genet Syst,2014,89(1):27-34
    [11]CHRISTIAENS O.Key role and diversity of EcR/USP and other nuclear receptors in selected Arthropoda species[D].Ghent:Ghent University,2013
    [12]ПРОХОРАМИ,ТУМАНОАСЮ,顾焕斌.动物体内胆固醇的生物合成[J].生理科学进展,1965,7(2):76-87
    [13]CHANNON H J.Cholesterol synthesis in the animal body[J].Biochem J,1925,19(3):424-432
    [14]JOUNI Z E,ZAMORA J,WELLS M A.Absorption and tissue distribution of cholesterol in Manduca sexta[J].Arch Insect Biochem Physiol,2002,49(3):167-175
    [15]ZDOBNOV E M,VON M C,LETUNIC I,et al.Comparative genome and proteome analysis of Anopheles gambiae and Drosophila melanogaster[J].Science,2002,298(5591):149-159
    [16]BEUMER H,LEHMANN F.ber die Cholesterinbildung im Tierk9rper[J].Z Gesamte Exp Med,1923,37(1):274-280
    [17]GILBERT L I,RYBCZYNSKI R,WARREN J T.Control and biochemical nature of the ecdysteroidogenic pathway[J].Annu Rev Entomol,2001,47:883-916
    [18]JOSHI M,AGARWAL H C.Site of cholesterol absorption in some insects[J].J Insect Physiol,1977,23(3):403-404
    [19]DYER D H,LOVELL S,THODEN J B,et al.The structural determination of an insect sterol carrier protein-2 with a ligand-bound C16 fatty acid at 1.35-resolution[J].J Biol Chem,2003,278(40):39085-39091
    [20]ENYA S,AMEKU T,IGARASHI F,et al.A Halloween gene noppera-bo encodes a glutathione S-transferase essential for ecdysteroid biosynthesis via regulating the behaviour of cholesterol in Drosophila[J/OL].Sci Rep,2014,4:6586[2018-03-25].https://www.ncbi.nlm.nih.gov/pubmed/25300303.DOI:10.1038/srep06586
    [21]WU F,WANG P,ZHAO Q,et al.Mutation of a cuticle protein gene,BmCPG10,is responsible for silkworm non-moulting in the2nd instar mutant[J/OL].PLoS ONE,2016,11(4):e0153549[2018-03-25].https://www.ncbi.nlm.nih.gov/pubmed/27096617.DOI:10.1371/journal.pone.0153549
    [22]WANG P,BI S,WU F,et al.Differentially expressed genes in the head of the 2nd instar pre-molting larvae of the nm2 mutant of the silkworm,Bombyx mori[J/OL].PLoS ONE,2017,12(7):e0180160[2018-03-25].https://www.ncbi.nlm.nih.gov/pubmed/28727825.DOI:10.1371/journal.pone.0180160
    [23]YOSHIYAMA T,NAMIKI T,MITA K,et al.Neverland is an evolutionally conserved rieske-domain protein that is essential for ecdysone synthesis and insect growth[J].Development,2006,133(13):2565-2574
    [24]THUMMEL C S,CHORY J.Steroid signaling in plants and insects:common themes,different pathways[J].Genes Dev,2002,16(24):3113-3129
    [25]WARREN J T,PETRYK A,MARQUES G,et al.Phantom encodes the 25-hydroxylase of Drosophila melanogaster and Bombyx mori:a P450 enzyme critical in ecdysone biosynthesis[J].Insect Biochem Mol Biol,2004,34(9):991-1010
    [26]WARREN J T,PETRYK A,MARQUES G,et al.Molecular and biochemical characterization of two P450 enzymes in the ecdysteroidogenic pathway of Drosophila melanogaster[J].Proc Natl Acad Sci USA,2002,99(17):11043-11048
    [27]PETRYK A,WARREN J T,MARQU S G,et al.Shade is the Drosophila P450 enzyme that mediates the hydroxylation of ecdysone to the steroid insect molting hormone 20-hydroxyecdysone[J].Proc Natl Acad Sci USA,2003,100(24):13773-13778
    [28]CHAVEZ V M,MARQU S G,DELBECQUE J P,et al.The Drosophila disembodied gene controls late embryonic morphogenesis and codes for a cytochrome P450 enzyme that regulates embryonic ecdysone levels[J].Development,2000,127(19):4115-4126
    [29]NAMIKI T,NIWA R,SAKUDOH T,et al.Cytochrome P450CYP307A1/Spook:a regulator for ecdysone synthesis in insects[J].Biochem Biophys Res Commun,2005,337(1):367-374
    [30]ONO H,REWITZ K F,SHINODA T,et al.Spook and Spookier code for stage-specific components of the ecdysone biosynthetic pathway in Diptera[J].Dev Biol,2006,298(2):555-570
    [31]URYU O,OU Q,KOMURA-KAWA T,et al.Cooperative control of ecdysone biosynthesis in Drosophila by transcription factors seance,ouija board,and molting defective[J].Genetics,2018,208(2):605-622
    [32]OU Q,MAGICO A,KINGJONES K.Nuclear receptor DHR4controls the timing of steroid hormone pulses during Drosophila development[J/OL].PLoS Biol,2011,9(9):e1001160[2018-03-25].https://www.ncbi.nlm.nih.gov/pubmed/21980261.DOI:10.1371/journal.pbio.1001160
    [33]ARBEITMAN M N,HOGNESS D S.Molecular chaperones activate the Drosophila ecdysone receptor,an RXR heterodimer[J].Cell,2000,101(1):67-77
    [34]KINGJONES K,THUMMEL C S.Nuclear receptors-a perspective from Drosophila[J].Nat Rev Genet,2005,6(4):311-323
    [35]ASHBURNER M.Sequential gene activation by ecdysone in polytene chromosomes of Drosophila melanogaster:Ⅱ.The effects of inhibitors of protein synthesis[J].Dev Biol,1974,39(1):141-157
    [36]ASHBURNER M.Sequential gene activation by ecdysone in polytene chromosomes of Drosophila melanogaster:Ⅰ.Dependence upon ecdysone concentration[J].Dev Biol,1973,35(1):47-61
    [37]BONNETON F,CHAUMOT A,LAUDET V.Annotation of Tribolium nuclear receptors reveals an increase in evolutionary rate of a network controlling the ecdysone cascade[J].Insect Biochem Mol Biol,2008,38(4):416-429
    [38]KOSTROUCHOVA M,KRAUSE M,KOSTROUCH Z,et al.Nuclear hormone receptor CHR3 is a critical regulator of all four larval molts of the nematode Caenorhabditis elegans[J].Proc Natl Acad Sci USA,2001,98(13):7360-7365
    [39]ZHAO X,QIN Z,LIU W,et al.Nuclear receptor HR3 controls locust molt by regulating chitin synthesis and degradation genes of Locusta migratoria[J].Insect Biochem Mol Biol,2017,92:1-11
    [40]GUO W C,LIU X P,FU K Y,et al.Functions of nuclear receptor HR3 during larval-pupal molting in Leptinotarsa decemlineata(Say)revealed by in vivo RNA interference[J].Insect Biochem Mol Biol,2015,63:23-33
    [41]ROBINSONRECHAVI M,GARCIA H E,LAUDET V.The nuclear receptor superfamily[J].J Cell Sci,2003,116(4):585-586
    [42]KINGJONES K,CHARLES J P,LAM G,et al.The ecdysoneinduced DHR4 orphan nuclear receptor coordinates growth and maturation in Drosophila[J].Cell,2005,121(5):773-784
    [43]SUTHERLAND J D,KOZLOVA T,TZERTZINIS G,et al.Drosophila hormone receptor 38:a second partner for Drosophila USP suggests an unexpected role for nuclear receptors of the nerve growth factor-induced protein B type[J].Proc Natl Acad Sci USA,1995,92(17):7966-7970
    [44]GU S H,HSIEH Y C,LIN P L.Stimulation of orphan nuclear receptor HR38 gene expression by PTTH in prothoracic glands of the silkworm,Bombyx mori[J].J Insect Physiol,2016,90:8-16
    [45]KOZLOVA T,LAM G,THUMMEL C S.The Drosophila DHR38 nuclear receptor is required for adult cuticle integrity at eclosion[J].Dev Dyn,2009,238(3):701-707
    [46]RUAUD A F,LAM G,THUMMEL C S.The Drosophila NR4A nuclear receptor DHR38 regulates carbohydrate metabolism and glycogen storage[J].Mol Endocrinol,2010,25(1):83-91
    [47]ALLEN A K,SPRADLING A C.The Sf1-related nuclear hormone receptor Hr39 regulates Drosophila female reproductive tract development and function[J].Development,2008,135(2):311-321
    [48]KAUR S,JOBLING S,JONES C S,et al.The nuclear receptors of Biomphalaria glabrata and Lottia gigantea:implications for developing new model organisms[J/OL].PLoS ONE,2015,10(4):e0121259[2018-04-25].https://www.ncbi.nlm.nih.gov/pubmed/25849443.DOI:10.1371/journal.pone.0121259
    [49]CRUZ J,NIEVA C,MAN PADR S D,et al.Nuclear receptor Bg FTZ-F1 regulates molting and the timing of ecdysteroid production during nymphal development in the hemimetabolous insect Blattella germanica[J].Dev Dyn,2008,237(11):3179-3191
    [50]YAMADA M,MURATA T,HIROSE S,et al.Temporally restricted expression of transcription factor beta FTZ-F1:significance for embryogenesis,molting and metamorphosis in Drosophila melanogaster[J].Development,2000,127(23):5083-5092
    [51]URNESS L D U,THUMMEL C S.Molecular analysis of a steroidinduced regulatory hierarchy:the Drosophila E74A protein directly regulates L71-6 transcription[J].EMBO J,1995,14(24):6239-6246
    [52]LING L,GE X,LI Z,et al.MicroRNA Let-7 regulates molting and metamorphosis in the silkworm,Bombyx mori[J].Insect Biochem Mol Biol,2014,53:13-21
    [53]CAI M J,ZHAO W L,JING Y P,et al.20-Hydroxyecdysone activates Forkhead box O to promote proteolysis during Helicoverpa armigera molting[J].Development,2016,143(6):1005-1015
    [54]FUTAHASHI R,OKAMOTO S,KAWASAKI H,et al.Genome-wide identification of cuticular protein genes in the silkworm,Bombyx mori[J].Insect Biochem Mol Biol,2008,38(12):1138-1146
    [55]LIANG J,ZHANG L,XIANG Z,et al.Expression profile of cuticular genes of silkworm,Bombyx mori[J/OL].BMC Genomics,2010,11:173[2018-03-25].https://www.ncbi.nlm.nih.gov/pubmed/20226095.DOI:10.1186/1471-2164-11-173
    [56]SOARES M P M,SILVATORRES F A,ELIASNETO M,et al.Ecdysteroid-dependent expression of the tweedle and peroxidase genes during adult cuticle formation in the honey bee,Apis mellifera[J/OL].PLoSONE,2011,6(5):e20513[2018-03-25].https://www.ncbi.nlm.nih.gov/pubmed/21655217.DOI:10.1371/journal.pone.0020513
    [57]TEETS N M,DENLINGER D L.Surviving in a frozen desert:environmental stress physiology of terrestrial Antarctic arthropods[J].JExp Biol,2014,217(1):84-93
    [58]QIAO L,XIONG G,WANG R X,et al.Mutation of a cuticular protein,BmorCPR2,alters larval body shape and adaptability in silkworm,Bombyx mori[J].Genetics,2014,196(4):1103-1115
    [59]ASANO T,TAOKA M,SHINKAWA T.Identification of a cuticle protein with unique repeated motifs in the silkworm,Bombyx mori[J].Insect Biochem Mol Biol,2013,43(4):344-351
    [60]JASRAPURIA S,SPECHT C A,KRAMER K J,et al.Gene families of cuticular proteins analogous to peritrophins(CPAPs)in Tribolium castaneum have diverse functions[J/OL].PLoS ONE,2012,7(11):e49844[2018-03-25].https://www.ncbi.nlm.nih.gov/pubmed/23185457.DOI:10.1371/journal.pone.0049844
    [61]MUN S,NOH M Y,DITTMER N T,et al.Cuticular protein with a low complexity sequence becomes cross-linked during insect cuticle sclerotization and is required for the adult molt[J/OL].Sci Rep,2015,5:10484[2018-03-25].https://www.ncbi.nlm.nih.gov/pubmed/25994234.DOI:10.1038/srep10484
    [62]SONG T Q,YANG M L,WANG Y L,et al.Cuticular protein Lm T-wdl1 is involved in molt development of the migratory locust[J].Insect Sci,2016,23(4):520-530
    [63]MERZENDORFER H.Insect chitin synthases:a review[J].J Comp Physiol B,2006,176(1):1-15
    [64]MERZENDORFER H,ZIMOCH L.Chitin metabolism in insects:structure,function and regulation of chitin synthases and chitinases[J].J Exp Biol,2003,206(24):4393-4412
    [65]CANDY D J,KILBY B A.Studies on chitin synthesis in the desert locust[J].J Exp Biol,1962,39(1):129-140
    [66]FUKAMIZO T,KRAMER K J.Mechanism of chitin hydrolysis by the binary chitinase system in insect moulting fluid[J].Insect Biochem,1985,15(2):141-145
    [67]RONG S,LI D Q,ZHANG X Y,et al.RNA interference to reveal roles ofβ-N-acetylglucosaminidase gene during molting process in Locusta migratoria[J].Insect Sci,2013,20(1):109-119
    [68]ZHENG Y P,KRELL P J,DOUCET D,et al.Cloning,expression,and localization of a molt-relatedβ-N-acetylglucosaminidase in the spruce budworm,Choristoneura fumiferana[J].Arch Insect Biochem Physiol,2008,68(1):49-59
    [69]YANG M,WANG Y,JIANG F,et al.miR-71 and miR-263 jointly regulate target genes chitin synthase and chitinase to control locust molting[J/OL].PLoS Genet,2016,12(8):e1006257[2018-03-25].https://www.ncbi.nlm.nih.gov/pubmed/27532544.DOI:10.1371/journal.pgen.1006257
    [70]GORMAN M J,ARAKANE Y.Tyrosine hydroxylase is required for cuticle sclerotization and pigmentation in Tribolium castaneum[J].Insect Biochem Mol Biol,2010,40(3):267-273
    [71]WITTKOPP P J,TRUE J R,CARROLL S B.Reciprocal functions of the Drosophila yellow and ebony proteins in the development and evolution of pigment patterns[J].Development,2002,129(8):1849-1858
    [72]BRODBECK D,AMHERD R,CALLAERTS P,et al.Molecular and biochemical characterization of the aaNAT1(Dat)locus in Drosophila melanogaster:differential expression of two gene products[J].DNA Cell Biol,1998,17(7):621-633
    [73]WALTER M F,BLACK B C,AFSHAR G,et al.Temporal and spatial expression of the yellow gene in correlation with cuticle formation and dopa decarboxylase activity in Drosophila development[J].Dev Biol,1991,147(1):32-45
    [74]WITTKOPP P J,VACCARO K,CARROLL S B.Evolution of yellow gene regulation and pigmentation in Drosophila[J].Curr Biol,2002,12(18):1547-1556
    [75]HAN Q,FANG J,DING H,et al.Identification of Drosophila melanogaster yellow-f and yellow-f2 proteins as dopachrome-conversion enzymes[J].Biochem J,2002,368(Pt 1):333-340
    [76]WITTKOPP P J,WILLIAMS B L,SELEGUE J E,et al.Drosophila pigmentation evolution:divergent genotypes underlying convergent phenotypes[J].Proc Natl Acad Sci USA,2003,100(4):1808-1813
    [77]ARAKANE Y,LOMAKIN J,BEEMAN R W,et al.Molecular and functional analyses of amino acid decarboxylases involved in cuticle tanning in Tribolium castaneum[J].J Biol Chem,2009,284(24):16584-16594
    [78]HAYASHI Y.Properties of xanthine dehydrogenase in the silkworm,Bombyx mori L[J].Nature,1960,186(4730):1053-1054
    [79]PHILLIPS J P,FORREST H S.Ommochromes and pteridines[M]//ASHBURMER M,WRIGHT T R F.The genetics and biology of Drosophila.London:Academic Press,1980:216-217
    [80]COATES B S,HELLMICH R L,LEWIS L C.Two differentially expressed ommochrome-binding protein-like genes(obp1 and obp2)in larval fat body of the European corn borer,Ostrinia nubilalis[J].J Insect Sci,2005,5(19):1-12
    [81]WANG M X,LU Y,CAI Z Z,et al.Phenol oxidase is a necessary enzyme for the silkworm molting which is regulated by molting hormone[J].Mol Biol Rep,2013,40(5):3549-3555
    [82]WANG M X,CAI Z Z,LU Y,et al.Expression and functions of dopa decarboxylase in the silkworm,Bombyx mori was regulated by molting hormone[J].Mol Biol Rep,2013,40(6):4115-4122
    [83]TRUMAN J W,RIDDIFORD L M.The morphostatic actions of juvenile hormone[J].Insect Biochem Mol Biol,2007,37(8):761-770
    [84]RIDDIFORD L M.Microarrays reveal discrete phases in juvenile hormone regulation of mosquito reproduction[J].Proc Natl Acad Sci USA,2013,110(24):9623-9624
    [85]杨文佳.桔小实蝇几丁质和保幼激素合成与代谢中重要基因的研究[D].重庆:西南大学,2014
    [86]ZAPATA R,PIULACHS M D,BELL S X.Inhibitors of 3-hydroxy-3-methylglutaryl-CoA reductase lower fecundity in the German cockroach:correlation between the effects on fecundity in vivo with the inhibition of enzymatic activity in embryo cells[J].Pest Manag Sci,2003,59(10):1111-1117
    [87]WANG Y,GUO B,ZHANG F,et al.Molecular cloning and functional analysis of the gene encoding 3-hydroxy-3-methylglutaryl coenzyme A reductase from hazel(Corylus avellana L.Gasaway)[J].J Biochem Mol Biol,2007,40(6):861-869
    [88]CABA J,BUESA C,HEGARDT F G,et al.Catalytic properties of recombinant 3-hydroxy-3-methylglutaryl coenzyme A synthase-1from Blattella germanica[J].Insect Biochem Mol Biol,1997,27(6):499-505
    [89]DAIMON T,KOZAKI T,NIWA R,et al.Precocious metamorphosis in the juvenile hormone-deficient mutant of the silkworm,Bombyx mori[J/OL].PLoS Genet,2012,8(3):e1002486[2018-03-25].https://www.ncbi.nlm.nih.gov/pubmed/22412378.DOI:10.1371/journal.pgen.1002486
    [90]HAMMOCK B D,SPARKS T C.A rapid assay for insect juvenile hormone esterase activity[J].Anal Biochem,1977,82(2):573-579
    [91]SHARE M R,ROE R M.A partition assay for the simultaneous determination of insect juvenile hormone esterase and epoxide hydrolase activity[J].Anal Biochem,1988,169(1):81-88
    [92]MAXWELL R A,WELCH W H,HORODYSKI F M,et al.Juvenile hormone diol kinase.II.Sequencing,cloning,and molecular modeling of juvenile hormone-selective diol kinase from Manduca sexta[J].J Biol Chem,2002,277(24):21882-21890
    [93]BONNING B C,WARD V K,VAN MEER M M,et al.Disruption of lysosomal targeting is associated with insecticidal potency of juvenile hormone esterase[J].Proc Natl Acad Sci USA,1997,94(12):6007-6012
    [94]LIU S,YANG B,GU J,et al.Molecular cloning and characterization of a juvenile hormone esterase gene from brown planthopper,Nilaparvata lugens[J].J Insect Physiol,2008,54(12):1495-1502
    [95]HALARNKAR P P,SCHOOLEY D A.Reversed-phase liquid chromatographic separation of juvenile hormone and its metabolites,and its application for an in vivo juvenile hormone catabolism study in Manduca sexta[J].Anal Biochem,1990,188(2):394-397
    [96]LASSITER M T,APPERSON C S,ROE R M.Juvenile hormone metabolism during the fourth stadium and pupal stage of the southern house mosquito,Culex quinquefasciatus Say[J].J Insect Physiol,1995,41(10):869-876
    [97]LI S,ZHANG Q R,XU W H,et al.Juvenile hormone diol kinase,a calcium-binding protein with kinase activity,from the silkworm,Bombyx mori[J].Insect Biochem Mol Biol,2005,35(11):1235-1248
    [98]ASHOK M,TURNER C,WILSON T G.Insect juvenile hormone resistance gene homology with the b HLH-PAS family of transcriptional regulators[J].Proc Natl Acad Sci USA,1998,95(6):2761-2766
    [99]WILSON T G,FABIAN J.A Drosophila melanogaster mutant resistant to a chemical analog of juvenile hormone[J].Dev Biol,1986,118(1):190-201
    [100]GUO E,HE Q,LIU S,et al.Met is required for the maximal action of20-Hydroxyecdysone during Bombyx metamorphosis[J/OL].PLoSONE,2012,7(12):e53256[2018-03-25].https://www.ncbi.nlm.nih.gov/pubmed/23300902.DOI:10.1371/journal.pone.0053256
    [101]KAYUKAWA T,MINAKUCHI C,NAMIKI T,et al.Transcriptional regulation of juvenile hormone-mediated induction of Krüppel homolog 1,a repressor of insect metamorphosis[J].Proc Natl Acad Sci USA,2012,109(29):11729-11734
    [102]DAIMON T,UCHIBORI M,NAKAO H,et al.Knockout silkworms reveal a dispensable role for juvenile hormones in holometabolous life cycle[J].Proc Natl Acad Sci USA,2015,112(31):4226-4235
    [103]KONOPOVA B,JINDRA M.Juvenile hormone resistance gene Methoprene-tolerant controls entry into metamorphosis in the beetle Tribolium castaneum[J].Proc Natl Acad Sci USA,2007,104(25):10488-10493
    [104]JESUS L,XAVIER B.Conserved repressive function of Krüppel homolog 1 on insect metamorphosis in hemimetabolous and holometabolous species[J/OL].Sci Rep,2011,1:163[2018-03-25].https://www.ncbi.nlm.nih.gov/pubmed/22355678.DOI:10.1038/srep00163
    [105]KONOPOVA B,SMYKAL V,JINDRA M.Common and distinct roles of juvenile hormone signaling genes in metamorphosis of holometabolous and hemimetabolous insects[J/OL].PLoS ONE,2011,6(12):e28728[2018-03-25].https://www.ncbi.nlm.nih.gov/pubmed/22174880.DOI:10.1371/journal.pone.0028728
    [106]WU B,MA L,ZHANG E,et al.Sexual dimorphism of sleep regulated by juvenile hormone signaling in Drosophila[J/OL].PLoS Genet,2018,14(4):e1007318[2018-03-25].https://www.ncbi.nlm.nih.gov/pubmed/29617359.DOI:10.1371/journal.pgen.1007318
    [107]KONOPOVA B,JINDRA M.Broad-Complex acts downstream of Met in juvenile hormone signaling to coordinate primitive holometabolan metamorphosis[J].Development,2008,135(3):559-568
    [108]ZHOU B,RIDDIFORD L M.Hormonal regulation and patterning of the broad-complex in the epidermis and wing discs of the tobacco hornworm,Manduca sexta[J].Dev Biol,2001,231(1):125-137
    [109]CAKOUROS D,DAISH T,MARTIN D,et al.Ecdysone-induced expression of the caspase DRONC during hormone-dependent programmed cell death in Drosophila is regulated by Broad-Complex[J].J Cell Biol,2002,157(6):985-995
    [110]JIANG C,LAMBLIN A F,STELLER H,et al.A steroid-triggered transcriptional hierarchy controls salivary gland cell death during Drosophila metamorphosis[J].Mol Cell,2000,5(3):445-455
    [111]RIDDIFORD L M,HIRUMA K,ZHOU X,et al.Insights into the molecular basis of the hormonal control of molting and metamorphosis from Manduca sexta and Drosophila melanogaster[J].Insect Biochem Mol Biol,2004,33(12):1327-1338
    [112]BAEHRECKE E H,THUMMEL C S.The Drosophila E93 gene from the 93F early puff displays stage-and tissue-specific regulation by 20-hydroxyecdysone[J].Dev Biol,1995,171(1):85-97
    [113]BELLES X,SANTOS C G.The MEKRE93(Methoprene tolerantKrüppel homolog 1-E93)pathway in the regulation of insect metamorphosis,and the homology of the pupal stage[J].Insect Biochem Mol Biol,2014,52(1):60-68
    [114]KAYUKAWA T,JOURAKU A,ITO Y,et al.Molecular mechanism underlying juvenile hormone-mediated repression of precocious larval-adult metamorphosis[J].Proc Natl Acad Sci USA,2017,114(5):1057-1062
    [115]NASSEL D R.Neuropeptides in the nervous system of Drosophila and other insects:multiple roles as neuromodulators and neurohormones[J].Prog Neurobiol,2002,68(1):1-84
    [116]MCBRAYER Z,ONO H,SHIMELL M,et al.Prothoracicotropic hormone regulates developmental timing and body size in Drosophila[J].Dev Cell,2007,13(6):857-871
    [117]MIZOGUCHI A,DEDOS S G,FUGO H,et al.Basic pattern of fluctuation in hemolymph PTTH titers during larval-pupal and pupaladult development of the silkworm,Bombyx mori[J].Gen Comp Endocrinol,2002,127(2):181-189
    [118]MIZOGUCHI A,OHASHI Y,HOSODA K,et al.Developmental profile of the changes in the prothoracicotropic hormone titer in hemolymph of the silkworm Bombyx mori:correlation with ecdysteroid secretion[J].Insect Biochem Mol Biol,2001,31(4/5):349-358
    [119]SHIOMI K,FUJIWARA Y,ATSUMI T,et al.Myocyte enhancer factor 2(MEF2)is a key modulator of the expression of the prothoracicotropic hormone gene in the silkworm,Bombyx mori[J].FEBS J,2005,272(15):3853-3862
    [120]SHIMELL M,PAN X,MARTIN F A,et al.Prothoracicotropic hormone modulates environmental adaptive plasticity through the control of developmental timing[J/OL].Development,2018,145(6):dev.159699[2018-03-25].https://www.ncbi.nlm.nih.gov/pubmed/29467242.DOI:10.1242/dev.159699
    [121]UCHIBORI-ASANO M,KAYUKAWA T,SEZUTSU H,et al.Severe developmental timing defects in the prothoracicotropic hormone(PTTH)-deficient silkworm,Bombyx mori[J].Insect Biochem Mol Biol,2017,87:14-25
    [122]RYBCZYNSKI R,BELL S C,GILBERT L I.Activation of an extracellular signal-regulated kinase(ERK)by the insect prothoracicotropic hormone[J].Mol Cell Endocrinol,2001,184(1/2):1-11
    [123]RYBCZYNSKI R,GILBERT L I.Prothoracicotropic hormone stimulated extracellular signal-regulated kinase(ERK)activity:the changing roles of Ca2+-and c AMP-dependent mechanisms in the insect prothoracic glands during metamorphosis[J].Mol Cell Endocrinol,2003,205(1):159-168
    [124]LIN J L,GU S H.In vitro and in vivo stimulation of extracellular signal-regulated kinase(ERK)by the prothoracicotropic hormone in prothoracic gland cells and its developmental regulation in the silkworm,Bombyx mori[J].J Insect Physiol,2007,53(6):622-631
    [125]GU S H,LIN J L,LIN P L.PTTH-stimulated ERK phosphorylation in prothoracic glands of the silkworm,Bombyx mori:role of Ca2+/calmodulin and receptor tyrosine kinase[J].J Insect Physiol,2010,56(1):93-101
    [126]CALDWELL P E,WALKIEWICZ M,STERN M.Ras activity in the Drosophila prothoracic gland regulates body size and developmental rate via ecdysone release[J].Curr Biol,2005,15(20):1785-1795
    [127]GU S H,YEH W L,YOUNG S C,et al.TOR signaling is involved in PTTH-stimulated ecdysteroidogenesis by prothoracic glands in the silkworm,Bombyx mori[J].Insect Biochem Mol Biol,2012,42(4):296-303
    [128]LAYALLE S,ARQUIER N,L OPOLD P.The TOR pathway couples nutrition and developmental timing in Drosophila[J].Dev Cell,2008,15(4):568-577
    [129]SCIEUZO C,NARDIELLO M,SALVIA R,et al.Ecdysteroidogenesis and development in Heliothis virescens(Lepidoptera:Noctuidae):focus on PTTH-stimulated pathways[J].J Insect Physiol,2018,107:57-67
    [130]REWITZ K F,YAMANAKA N,GILBERT L I,et al.The insect neuropeptide PTTH activates receptor tyrosine kinase torso to initiate metamorphosis[J].Science,2009,326(5958):1403-1405
    [131]ZHANG M,XU W H.Isolation of an eclosion hormone gene from the cotton bollworm,Helicoverpa armigera:temporal and spatial distribution of transcripts[J].Comp Biochem Physiol,B,2006,143(3):351-359
    [132]于淼.家蚕PTTH基因转录调控及相关神经肽的发育表达[D].合肥:合肥工业大学,2010
    [133]ITNˇAN D,ADAMS M E.Excitatory and inhibitory roles of central ganglia in initiation of the insect ecdysis behavioural sequence[J].J Exp Biol,2000,203(8):1329-1340
    [134]ABDEL-LATIEF M,MEYERING-VOS M,HOFFMANN K H.TypeA allatostatins from the fall armyworm,Spodoptera frugiperda:molecular cloning,expression and tissue-specific localization[J].Arch Insect Biochem Physiol,2004,56(3):120-132
    [135]SUI Y P,LIU X B,CHAI L Q,et al.Characterization and influences of classical insect hormones on the expression profiles of a molting carboxypeptidase A from the cotton bollworm(Helicoverpa armigera)[J].Insect Mol Biol,2009,18(3):353-363
    [136]RUSSEL S,FRAND A R,RUVKUN G.Regulation of the C.elegans molt by pqn-47[J].Dev Biol,2011,360(2):297-309
    [137]SATO K,MIYATA K,OZAWA S,et al.Systemic RNAi of V-ATPase subunit B causes molting defect and developmental abnormalities in Periplaneta fuliginosa[J/OL].Insect Sci,2017[2018-03-25].https://www.ncbi.nlm.nih.gov/pubmed/29285882.DOI:10.1111/1744-7917.12565
    [138]MANSUR J F,ALVARENGA E S,FIGUEIRA-MANSUR J,et al.Effects of chitin synthase double-stranded RNA on molting and oogenesis in the Chagas disease vector Rhodnius prolixus[J].Insect Biochem Mol Biol,2014,51:110-121
    [139]BROEHAN G,ARAKANE Y,BEEMAN R W,et al.Chymotrypsinlike peptidases from Tribolium castaneum:a role in molting revealed by RNA interference[J].Insect Biochem Mol Biol,2010,40(3):274-283
    [140]ARAKANE Y,BAGUINON M C,JASRAPURIA S,et al.Both UDP N-acetylglucosamine pyrophosphorylases of Tribolium castaneum are critical for molting,survival and fecundity[J].Insect Biochem Mol Biol,2011,41(1):42-50
    [141]ZHANG J,LU A,KONG L,et al.Functional analysis of insect molting fluid proteins on the protection and regulation of ecdysis[J].J Biol Chem,2014,289(52):35891-35906