拟黑多刺蚁semaphorin 2a基因的克隆与mRNA水平表达的研究
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摘要
拟黑多刺蚁(Polyrhachis vicina Roger)隶属于膜翅目(Hymenoptera),蚁科(Formicidae),多刺蚁属(Polyrhachis),是一类营群居的典型的社会性昆虫,同时也是一类重要的资源昆虫。该种昆虫具有独特的品级分化、行为复杂、分工明确等社会性特征以及高度复杂的神经系统,因此是研究昆虫发育、行为调控机制及神经功能方面的很好材料。
     Semaphoring 2a(sema2a)是semaphorin家族中的第二亚家族中的的分泌蛋白,曾经在果蝇(Drosophila melanogaster Meigen)、沙漠蝗虫(Schistocerca gregaria Forskal)、蟋蟀(Gryllusbimaculatus De Geer)中发现,该蛋白可以作为选择性靶标抑制突触的形成。
     本论文以拟黑多刺蚁为研究材料。通过RT-PCR及RACE技术从拟黑多刺蚁体内克隆sema2a基因的全长cDNA序列,并对其核苷酸序列利蛋白序列进行生物信息学分析;通过实时定量RT-PCR方法对sema2a基因在拟黑多刺蚁体内的mRNA表达水平进行相对定量研究;研究结果如下:
     1.拟黑多刺蚁sema2a基因cDNA序列全长2763 bp,包含一个长度为2151 bp的开放阅读框,编码716个氨基酸残基。5'端非编码区(5'-UTR)长度为92 bp,3'-UTR为521 bp,3'-UTR区可见典型的加尾信号AATAAA及18bp长度的PolyA尾。编码的蛋白质分子量为81.1 kDa,等电点pI=7.05。同源性分析表明该蛋白与意大利蜜蜂mAchR的氨基酸序列相似性达98%,与金小蜂、拟谷盗、沙漠蝗虫和果蝇的氨基酸序列相似性分别为98%、95%、83%、71%和69%。生物信息学分析表明,该蛋白为分泌型蛋白,这一结论与报道一致。因此,可以确定本次克隆所获得序列为拟黑多刺蚁sema2a基因的全长cDNA序列,命名为Pv-sema-2a,上传GenBank后,获得的基因序列号为EU836737。
     2.采用荧光实时定量RT-PCR方法对拟黑多刺蚁各发育阶段虫体、不同品级成虫及不同品级成虫头部的Pv-sema-2a mRNA表达水平进行相对定量分析。结果表明,Pv-sema-2a基因在所检测的各个样本内均有表达,从整个发育阶段来说,从胚胎到第一龄幼虫,Pv-sema-2a的表达量有所增加,但随着幼虫的发育,Pv-sema-2a的表达量逐渐降低,直到蛹期,其表达量又再次升高。在三个品级成虫比较中,各品级之间的表达量有明显的不同,工蚁的表达量最高,雌蚁次之,在雄蚁成体中的表达量最低。在三个品级成虫头部比较中,雌蚁头中的表达量最高,其表达量为工蚁头的1.25倍,雄蚁头的表达量为工蚁头的0.26倍。Pv-sema-2a基因在胚胎和蛹中的高表达显示了该基因在蚂蚁发育中的作用。
     3.采用荧光实时定量RT-PCR方法对拟黑多刺蚁行为学训练后,已训练的拟黑多刺蚁和未训练的拟黑多刺蚁之间头部Pv-sema-2a mRNA表达水平进行相对定量分析。发现已训练的拟黑多刺蚁和未训练的拟黑多刺蚁之间头部Pv-sema-2a mRNA表达差异显著,未训练的拟黑多刺蚁头部的表达要比训练过的拟黑多刺蚁头部表达量高。由此可以推出Pv-sema-2a mRNA在经过训练的拟黑多刺蚁头部低表达可能有助于新的突触连接的建立,与增强学习和记忆有关。
     本项研究首次从社会性昆虫拟黑多刺蚁中克隆sema基因的全长cDNA序列,并将全长cDNA序列上传GenBank;采用相对实时定量方法研究发现,Pv-sema-2a mRNA在拟黑多刺蚁不通不同发育阶段虫体、不同品级成虫及不同品级成虫头部差异表达:并对成年拟黑多刺蚁工蚁进行行为训练,用相对实时定量的方法发现,未训练的拟黑多刺蚁头部的表达要比训练过的拟黑多刺蚁头部表达量高。这些研究结果将为深入探讨昆虫sema2a的相关研究奠定基础。
Polyrhachis vicina Roger as eusocial insects belongs to the genus Polyrhachis(Hymenoptera: Formicidae).This species possess the characteristic of castes differentiation,sophisticated behaviors, behavioral plasticity and highly complex nervous system.At the same time,p.vicina is also a very important resource insect.Consequently,P.vicina as a good material is used for study the mechanisms of insect development,behavior,and the function of insects'nerve system.
     Semaphorin 2a belongs to the secreted members of the semaphorin family identified as class 2 and only present in invertebrates,which has been identified in the fruit fly(Drosophila melanogaster Meigen),grasshopper(Schistocerca gregaria Forskal),and Cricket(Gryllus bimaculatus De Geer). Semaphorin 2a may be acting the selective target to inhibiting synaptic arborization formation.
     In this paper,the full-length cDNA of sema2a gene is coloned from the P.vicina by RT-PCR and RACE methods.The bioinformatics method is used to analyse characteristics of full-length cDNA and predict functional motifs in the ORF.The mRNA expression levels of sema2a in the ant were investigated by real-time RT-PCR.The major experiment results are as follow:
     1.The full-length cDNA of Pv-sema-2a is 2763 bp,including an open reading frame(ORF) of 2151 bp,which encodes a deduced 599-amino acid peptide with a predicted molecular mass of 81.1 kDa and with the theoretical pI of 7.05,which contains a 5'-untraslated region(5'-UTR) of 92 bp and a 3'-UTR of 521 bp,and a putative polyadenylation signal AATAAA was found at 18 bp upstream from the 16-nucleotide poly(A) tail in 3'-UTR.The results of sequence alignments analysis showed that full length cDNA of Pv-sema-2a is similar to Apis mellifera,Nasonia vitripennis,Tribolium castaneum,Schistocerca gregaria and Drosophila melanogaster at 98%,95%,83%,71%and 69%. From the results of bioinformation analysis show that this protein is a secreted member,this is identical with previous research.The nucleotide sequence of Pv-sema-2a gene is submitted to GenBank and assigned the number EU836737.
     2.The relative quantification expression of the mRNA level of Pv-sema-2a genes in different developmental periods and the different castes were analyzed using real-time quantitative RT-PCR. The results showed that Pv-sema-2a mRNA is expressed in all samples tested.During development, the expression of Pv-sema-2a in first instar higher than embryos,and then declined gradually from the second instar to the fourth instar,but in pupa expression increased again,In three castes,the Pv-sema-2a expression was higest in worker ants,then female ants,lowest in male.The Pv-sema-2a mRNA expression in the heads of different castes from height to low was female,worker and male ants.The high mRNA levels in both the embryo and pupa suggest the essential role of Pv-sema-2a in ant development.
     3.To obtain information on whether sema2a associated with learning and memory,we analyzed Pv-sema-2a expression levels from the trained ants and untrained ants.We found Pv-sema-2a was expressed at a greatly lower in the trained ants than that in untrained ants.Consequently,the low expression of Pv-sema-2a may be beneficial in establishing these new connections.
     For the first time,we have cloned the full-length cDNAs of Pv-sema-2a genes from the eusocial insect,P.vicina,and submitted the cDNA sequences to GenBank.Real-time quantitative RT-PCR analysis revealed that Pv-sema-2a express at distinct developmental stages,different castes,heads and between the trained and untrained ants.These results may provide the theory foundation for further researching on the concrete function of Pv-sema-2a in insects.
引文
[1]唐觉等编著.中国经济昆虫志[M],第四十七册,膜翅目,蚁科.北京:科学出版社,1995.
    [2]贺传生,余华星.拟黑多刺蚁生活习性的初步研究[J].白蚁科技,1994,11(3):32-33
    [3]杨沛.蚁群社会生物学及多样性[J].松辽学刊(自然科学版),1999,36(4):21-24.
    [4]翟新国.“拟黑多刺蚁”蚂蚁生物学特性观察[J].邯郸农业高等专科学校学报,2000,17(4):11-12.
    [5]段广勋,王军华,王平.要用蚂蚁的研究进展[J]_微量元素与健康,1998,15(1):51。
    [6]杨大荣,舒畅,李朝达.五种昆虫的营养成分分析[J].营养学报,1996,18(2):231-234.
    [7]陈颖,陈德锋,汪树理.拟黑多刺蚁氨基酸和营养元素的分析[J].长春中医药大学学报,2008,24(3):257-258.
    [8]王荫长.昆虫生理生化学[M].北京:中国农业出版社,1994:116-139.
    [9]赵一,陈文.蚂蚁的药用价值[J].广西中医药,1989:12(2):35.
    [10]U.Yazdani,J.R.Terman,The semaphorins[J].Genome Biology,2006,7(3):211.
    [11]A.L.Kolodkin,D.J.Matthes,and C.S.Goodman.The semaphoring Genes Encode a Family of Transmembrane and Secreted Growth Cone Guidance Molecules[J].Cell,1993,75(7):1389-1399.
    [12]Unified nomenclature for the semaphorins/collapsins[J].Semaphorin Nomenclature Committee.Cell,1999,97:551-552.
    [13]D.J.Matthes,H.Sink,A.L.Kolodkin,and C.S.Goodman.Semaphorin Ⅱ Can Function as a Selective Inhibitor of Specific Synaptic Arborizations[J].GoodmanCell.1995,81(4):631-639.
    [14]K.E.Bates,P.M.Whitington.Semaphorin 2a secreted by oenocytes signals through plexin B and plexin A to guide sensory axons in the Drosophila embryo[J].Developmental Biology.2007,302(2):522-535.
    [15]J.C.Ayoob,J.R.Terman,and A.L.Kolodkin.Drosophila Plexin B is a Sema-2a receptor required for axon guidance[J].Development.2006,133(11):2125-2135.
    [16]P.J.Roy,H.Zheng,C.E.Warren and J.G.Culotti.mab-20 encodes Semaphorin-2a and is required to prevent ectopic cell contacts during epidermal morphogenesis in Caenorhabditis elegans[J].Development.2000,127(4):755-767.
    [17]I.D.Chin-Sang,L.S.Moseley,M.Ding,J.R.Harrington,S.E.George and A.D.Chisholm.The divergent C.elegans ephrin EFN-4 functions inembryonic morphogenesis in a pathway independent of the VAB-1 Eph receptor.Development,2002,129,5499-5510.
    [18]T.Fujii,E Nakao,Y.Shibata,G.Shioi,E.Kodama,H.Fujisawa,S.Takagi.Caenorhabditis elegans PlexinA,PLX-1,interacts with transmembrane semaphorins and regulates epidermal morphogenesis.Development,2002,129(9):2053-2063.
    [19]C.M.Isbister,A.Tsai,S.T.Wong,A.L.Kolodkin and T.P.O'Connor.Discrete roles for secreted and transmembrane semaphorins in neuronal growth cone guidance in vivo[J].Development.1999,126(9),2007-2019.
    [20]C.M.Isbister,P.J.Mackenzie,K.C.To,and T.P.O'Connor.Gradient Steepness Influences the Pathfinding Decisions of Neuronal Growth Cones In Vivo[J].The Journal of Neuroscience.2003,23(1):193-202.
    [21]K.R.Maynard,S.S.McCarthy,E.Sheldon,and H.W.Horch.Developmental and Adult Expression of Semaphorin 2a in the Cricket Gryllus bimaculatus[J].The Journal of Comparative Neurology.2007,503(1):169-181.
    [22]朱珊丽,尤孙武,娄崇洁,龚红君,张丽芳.沙眼衣原体主要外膜蛋白生物信息学分析[J].温州医学院学报,2009,39(1):5-7.
    [23]J.Chory,J.R.Ecker,Briggs S.National Science Foundation-Sponsored Workshop Report:"The 2010 Project" functional genomics and the virtual plant.A blueprint for understanding how plants are built and how to improve them[J].Plant Physiol,2000,123(2):423-426.
    [24]M.Y.Galperin.The Molecular Biology Database Collection:2005 update[J].Nucleic Acids Research,2005,33(Database issue):15-24.
    [25]D.Gibert.Bioinfamatics software resources[J].Brief Bioinform,2004,5(3):300-304.
    [26]J.Skolnick,J.S.Fetrow.From genes to protein structure and function:novel applications of computational approaches in genomic era[J].Trends in iotechnology,2000,18(1):34-39.
    [27]青木清,庚镇城.行为生物学[J].世界科学,1983,3:31-32.
    [28]孙忻 王丽.动物的本能行为和学习行为[J].动物之美,2000,3:30-31.
    [29]陈阅增.普通生物学[M].北京:高等教育出版社,1997:235.
    [30]R.J.Prokopy,A.H.Reynolds,L.J.Ent.Can Rhagoletis pomonella flies(Diptera:Tephritidae )learn to associate presence of food on foliage with leaf color[J].European Jounral of Entomolog,1998,95(3):335-341.
    [31]R.Dukas,E.A.Bernays.Learning improves growth rate in grasshoppers[C].Proceedings of the National Academy of Sciences of the United States of America,2000,97:2637-2640.
    [32]D.Raubenheimer,J.Blackshaw.Locusts learn to associate visual stimuli with drinking[J].J.Insect.Behav.,1994,7(4):569-575.
    [33]T.M.Alloway.Learning and memory in insects[J].Ann.Rev.Ent,1972,17:43-56.
    [34]H.Markl Insect behavior:functions and mechanisms[M].New York:Academic Press,1974:3-14
    [35]S.A.Bustin.Absolute quantification of mRNA using real-time reverse transcription Polymerase chain reaction assyas[J].J.Mol.Endoerinol.2000,25(2):169-193.
    [36]邓少丽,罗阳,陈伟.荧光定量PCR技术及其在临床上的应用[J].国外医学临床生物化学与检验学分册,2002,23(5):301-303.
    [37]杨龙.猪MEF2A基因的克隆及表达规律的研究[D].哈尔滨:东北农业大学,2008.
    [38]王礼坚.蚂蚁的养殖技术.中药材,17(8):10.
    [39]谭振.纸箱立体养蚂蚁技术.特种经济动植物.2001,05期:10.
    [40]J.Sambrook,E.F.Fritsch,T.Maniatis.Molecular Cloning:A Laboratory Manual (2nd Ed.)[M].Cold Spring Harbor University Press,Cold Spring Harbor,NY,1989.
    [41]F.M.Ausubel.Current Protocols in Molecular Biology[M].John Wiley and Sons,Inc.,NY,1999.
    [42]F.Jeanmougin,J.D.Thompson,M.Gouy,D.G.Higgins,T.J.Gibson.Multiple Sequence Alignment with Clustal X[J].Trends in Biochemical Sciences.1998,23(10):403-405.
    [43]任南琪,林海龙,李建政,郑国香,李永丰,于振国,张昆.用CODEHOP设计简并引物克隆B49乙酸激酶基因片段[J].哈尔滨工业大学学报,2007,39(8):1225-1229.
    [44]黄菁,王少丽,乔传令.程序化设计简并引物与克隆小菜蛾酯酶基因[J].昆虫 知识,2002,39(6):458-461.
    [45]夏瑞,陆旺金,李建国,杜娟.简并引物的程序化设计与荔枝HMGR基因片段的克隆[J].果树学报,2006,23(6):903-906.
    [46]M.Kozak.Interpreting cDNA Sequences;Some Insights from Studies on Translation[J].Mamm Genoma.1996,7(8):563-574.
    [47]M.Fitzgerald,and T.Shenk.The sequence 5'-AAUAAA-3'forms parts of the recognition site for polyadenylation of late SV40 mRNAs.Cell,1981,24(1):251-260.
    [48]张庆华,茅矛,陈竺.基因组研究中全长cDNA克隆的策略[J].生物工程进展,2000,20(4):3-5.
    [49]顾红雅,翟礼嘉等.植物基因与分子操作[M].北京:北京大学出版社,1995:74-931.
    [50]M.A.Frohman,M.K.Dush,G.R.Mrtin,et al.Rapid production of full -length cDNAs from rare transcripts:amplification using single gene specific oligonuleotide primer[J].Proc Natl Acad Sci USA,1988,85(23):8998 - 9002.
    [51]E.Gasteiger,C.Hoogland,A.Gattiker,et al.Protein Identification and Analysis Tools on the Expasy Server;(In) John M.Walker(Ed):The Proteomics Protocols Handbook[M].Humana Press,2005:571-607.
    [52]K.J.Jensen,S.M.Brunaks.Prediction of Novel Archaeal Enzymes from Sequence-Derived Features[J].Protein Science.2002,11(12):2894-2898.
    [53]J.D.Bendtsen,H.Nielsen,C.S.Vonheijneq Brunal.Improved Prediction of Signal Peptides:Signal P3.0[J].J Mol Biol.2004,340:783-795.
    [54]H.Nielsen,J.Engelbrecht,S.Brunak,et al.Identification of Prokaryotic and Eukotic Signal Peptides and Prediction of Their Cieavage Sites[J].Pro Eng.1997,10:1-6.
    [55]S.F.Altschul,L.M.Thomas,A.A.Schffer,et al.Gapped BLAST and PSI-BLAST:A New Generation of Protein Database Search Programs[J].Nucleic Acids Research.1997,25(17):3389-3402.
    [56]A.A.Schffer.L.Arabind,T.L.Madden,et al.Improving the Accuracy of PSI-BLAST Protein Database Searches with Composition-Based Statistics and Other Refinements[J].Nucleic Acids Research.2001,29(14):2994-3005.
    [57]T.Schwede,J.Kopp,N.Guex,M.C.Peitsch SWISS-MODEL:An Automated Protein Homology-Modeling Server[J].Nucleic Acids Research.2003,31(13): 3381-3385.
    [58]N.Guex,M.C.Peitsch.SWISS-MODEL and the Swiss-Pdbviewer:An Environment for Comparative Protein Modeling[J].Electrophoresis.1997,18(15):2714-2723.
    [59]K.Arnold,L.Bordoli,J.Kopp,T.Schwede.The SWISS-MODEL Workspace:A Web-Based Environment for Protein Structure Homology Modeling[J].Bioinformatics.2006,22(2):195-201.
    [60]K.Tamura,J.Dudley,M.Nei,S.Kumar.MEGA4:Molecular Evolutionary Genetics Analysis(MEGA) Software Version 4.0[J].Molecular Biology and Evolution.2007,24(8):1596-1599.
    [61]N.Saitou,M.Nei.The Neighbor-Joining Method:A New Method for Reconstructing Phylogenetic Trees[J].Molecular Biology and Evolution.1987,4(4):406-425.
    [62]D.Fischer,D.Eisenberg.Assigning Folds to the Proteins Encoded Bye the Genome of Mycoplasma Genitalium[J].Proceedings of the National Academy of Sciences USA.1997,94(28):11929-11934.
    [63]D.Xu,M.A.Unseren,Y.Xu,E.C.Uberbacher.Sequence-Structure Specificity of a Knowledge Based Energy Function at the Secondary Structure Level[J].Bioinformatics.2000,16(3):257-268.
    [64]N.Siew,D.Fischer.Convergent Evolutions of Protein Structure Prediction and Computer Chess Tournarnents:CASP,Kasparov and CAFASP[J].IBM Sys J.2001,40(2):410-425.
    [65]A.Sali.Modeling Muations and Homologous Proteins[J].Current Opinion in Biotechnology.1995,6(4):437-445.
    [66]M.C Peitsch.Protein modeling by e-mail.From amino acid sequence to protein structure:A free one-hour service[J].Nature Biotechnology 1995,7(13):658-660.
    [67]J.Skolnick,J.S.Fetrow.From genes to protein structure and function:Novel applications of computational approaches in genomic era[J].TIBTECH.2000,18(1):34-39.
    [68]王璐.拟南芥MPK 17基因的生物信息学分析及功能的初探[D].山西:山西大学,2008.
    [69]石嵘,马文丽,郑文岭.后基因组研究中蛋白结构与功能的预测[J].生物技术通报,2002,(3):1-4.
    [70]C.A.Love,K.Harlos,N.Mavaddat,S.J.Davis,D.I.Stuart,E.Y.Jones,R.M.Esnouf.The ligand-binding face of the semaphorins revealed by the high-resolution crystal structure of SEMA4D[J].Nat Struct Biol.2003,10(10):843-848.
    [71]欧阳霞辉.拟黑多刺蚁和意大利蜜蜂雌激素相关受体基因的克隆与表达研究[D].西安:陕西师范大学,2007.
    [72]Y.Ishii,K.Kubota,K.Hara.Postembryonic Development of the mushroom bodies in the ant,Camponotus Japonicus[J].Zoological Science.2005,22(7):743-753.
    [73]T.Suzuki,P.J.Higgins,D.R.Crawford.Control Selection for RNA Quantitation[J].Biotechniques.2000,29(2):332-337.
    [74]I.Claeys,G.Simonet,T.Van-Loy,et al.cDNA Cloning and Transcript Distribution of Two Novel Members of the Neuroparsin Family in the Desert Locust,Schistocerca Gregaria[J].Insect Molecular Biology.2003,12(5):473-481.
    [75]G.Simonet,I.Claeys,B.Breugelmans,et al.Transcript Profiling of Pacifastin-Like Peptide Precursors in Crowd and Isolated-Reared Desert Locusts[J].Biochemical and Biophysical Research Communications.2004,317(2):565-569.
    [76]张驰宇,徐顺高,黄新祥.一种新颖简便的荧光实时RT-PCR相对定量方法的建立[J].生物化学与生物物理学进展,2005,32(9):883-888.
    [77]S.A.Bustin.Absolute Quantification of mRNA Using Real-Time Reverse Transcription Polymerase Chain Reaction Assays[J].J.Mol.Endocrinol.2000,25(2):169-193.
    [78]Giulietti,L.Overgergh,D.Valckx.An Overview of Real-Time Quantitative PCR:Applications to Quantify Cytokine Gene Expression[J].Methods.2001,25(4):386-401.
    [79]K.J.Livak,T.D.Schmittgen.Analysis of Relative Gene Expression Data Using,Real-Time Quantitative PCR and the 2(-Delta Delta C(T)) Method[J].Methods.2001,25(4):402-408
    [80]M.Nemoto,,K.Hara.Ecdysone receptor expression in developing and adult mushroom bodies of the ant Camponotus japonicus[J].Development Genes and Evolution.2007,217(9):619-627.
    [81]S.E.Fahrbach.2006.Structure of the mushroom bodies of the insect brain[J].Annu Rev Entomol 51:209-232.
    [82]A.Pascual,T.Preat.Localization of long-term memory within the Drosophila mushroom body.Science.2001,294(5544):1115-1117.
    [83]M.Mizunami,J.M.Weibrecht,N.J.Strausfeld.Mushroom bodies of the cockroach:their participation in place memory.J Comp Neurol.1998,402(4):520-537.
    [84]L.Liu,R.Wolf,R.Ernst,M.Heisenberg.Context generalization in Drosophila visual learning requires the mushroom bodies.Nature.1999,400(4746):753-756.
    [85]R.Gerlai.A new continuous altemation task in T-maze detects hippocampal dysfunction in mice A strain comparison and lesion study[J].Behavioural Brain Research,1998,95(1):91-101.
    [86]金卓明.CREB在拟黑多刺蚁学习记忆中的作用研究[D].西安:陕西师范大学,2006.
    [87]王荫长.昆虫生理生化学[M].北京:中国农业出版社,1994,116-139.
    [88]黄明.噪声对小鼠学习和记忆的影响及其相关机制的研究[D].武汉:华中师范大学,2008.
    [89]柯尊记,王亚威,周兰仙等.大鼠乳头体核突触的生后发育-亚突触结构的定量研究[J]。神经解剖学杂志,1992,8:211
    [90]C.W.Cotman,and M.Niet-Sampedro.Cell biology of synaptic plasticity[J].Science,1984,225(4668):1287
    [91]D.Purves.Elimination of synapse in the developing nervous system.Science,1980,210(4466):153.
    [92]G.S.Withers,S.E.Fahrbach,G.E.Robinson.Selective neuroanatomical plasticity and division of labour in the honeybee.Nature,1993,364(6434):238-240.
    [93]W.Gronenberg,S.Heeren and B.Holldobler Age-dependent and task-related morphological changes in the brain and the mushroom bodies of the ant Camponotus floridanus[J].The journal of Experimental Biology,1996,199(9):2011-2019.
    [94]D.Yu,A.Ponomarev,R.L.Davis.Altered representation of the spatial code for odors after olfactory classical conditioning:memory trace formation by synaptic recruitment.Neuron,2004,42(3):437-449.
    [95]M.Cayre,C.Strambi,P.Charpin,R.Augier,M.R.Meyer,J.S.Edwards,A.Strambi.Neurogenesis in adult insect mushroom bodies[J].J Comp Neurol,1996,371(2):300-310.
    [96]S.E.Fahrbach,L.Strande,G.E.Robinson.Neurogenesis is absent in the brains of adult honey bees and does not explain behavioural neuroplasticity[J].Neurosci Lett, 1995, 197(2):145-148.
    [97]G. S. Withers, S. E. Fahrbach, G. E. Robinson. Selective neuroanatomical plasticity and division of labour in the honey bee [J]. Nature. 1993, 364: 238 -240.
    [98] J. S. Edwards. Postembryonic development and regeneration of the insect nervous system. Adv. Insect Physiol. 1969, 6: 97-137.
    [99] G. M. Technau. Fiber number in the mushroom bodies of adult Drosophila melanogaster depends on age, sex, and experience [J]. J. Neurogenet., 1984, 1(2):113-126.
    [100] M. L. Winberg, K. J. Mitchell, C. S. Goodman. Genetic Analysis of the Mechanisms Controlling Target Selection: Complementary and Combinatorial Functions of Netrins, Semaphorins, and IgCAMs [J]. Cell, 1998, 93(6): 581-591.

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