昆虫受精孔形态及其功能研究进展
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摘要
昆虫受精孔是在卵子发生后期,由滤泡细胞分化形成的微小孔道,在多数昆虫的卵壳上形成开口,是精子进入成熟卵的重要通道。受精孔区的形态结构或受精孔数目在不同昆虫中均存在较大差异,已作为重要的形态依据应用于一些种类的分类学研究中,同时研究表明鳞翅目昆虫受精孔数目与其生殖特性具有协同进化关系。近年来有研究发现,许多昆虫的受精孔数目常受交配等因素影响而发生变化,且在少数种类的突变体中也具有显著变化。目前,国内外关于受精孔的研究主要集中于受精孔区的形态观察与描述,而对其形态和数量变化的生物学功能及分子调节机制研究较少。故对昆虫受精孔的研究现状进行全面总结,并就未来的研究方向进行展望,以期为人们深入研究昆虫受精孔的形成机制及在害虫防治中的应用提供理论依据。
Micropyle is a small opening in egg shell that allows sperm to enter into the mature oocyte and they are deposited by the follicle cells during the late oogenesis phase. The morphology, structure, and number of micropyles are vastly different among different insect species. Thus, these features have been used as important identification characteristics in the taxonomic study of several insect species. This research conducted in Lepidoptera also revealed that the number variation of micropyles was co-evolved with their reproductive characteristics. In recent years, several studies have shown that the number of micropyles was changeable when the females were under different mating status(mate or virgin female) or paired with different males. In addition, for some mutants, their eggs had significantly different number of micropyles compared with those of the typical phenotype. In summary, the present studies were mostly focused on the morphology observation and description; while studies related to the regulation mechanisms of the number variation of micropyles and their biological functions were extreme scarce This paper has summarized the literature on various aspects of micropyles research and highlighted the perspectives of future studies. This paper maybe provide new insight for revealing the regulation mechanism of micropyles formation and the application prospect in pest management.
Micropyle is a small opening in egg shell that allows sperm to enter into the mature oocyte and they are deposited by the follicle cells during the late oogenesis phase. The morphology, structure, and number of micropyles are vastly different among different insect species. Thus, these features have been used as important identification characteristics in the taxonomic study of several insect species. This research conducted in Lepidoptera also revealed that the number variation of micropyles was co-evolved with their reproductive characteristics. In recent years, several studies have shown that the number of micropyles was changeable when the females were under different mating status(mate or virgin female) or paired with different males. In addition, for some mutants, their eggs had significantly different number of micropyles compared with those of the typical phenotype. In summary, the present studies were mostly focused on the morphology observation and description; while studies related to the regulation mechanisms of the number variation of micropyles and their biological functions were extreme scarce This paper has summarized the literature on various aspects of micropyles research and highlighted the perspectives of future studies. This paper maybe provide new insight for revealing the regulation mechanism of micropyles formation and the application prospect in pest management.
引文
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[35] KUEROVá Z,STEJSKAL V.External egg morphology of two stored product anobiids,Stegobium paniceum and Lasioderma serricorne(Coleoptera:Anobiidae)[J].Journal of Stored Products Research,2010,46(3):202-205.
[36] KU?EROVá Z,STEJSKAL V.Differences in egg morphology of the stored-grain pests Rhyzopertha dominica and Prostephanus truncatus(Coleoptera:Bostrichidae)[J].Journal of Stored Products Research,2008,44(1):103-105.
[37] VOGELGESANG M,SZKLARZEWICZ T.Formation and structure of egg capsules in scale insects (Hemiptera,Coccinea):I.Ortheziidae[J].Arthropod Structure & Development,2001,30(1):63-68.
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[43] GOBIN B,PEETERS C,BILLEN J.Production of trophic eggs by virgin workers in the ponerine ant Gnamptogenys menadensis[J].Physiological Entomology,1998,23(4):329-336.
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[2] CLARK J,LANGE A B.Evidence of a neural loop involved in controlling spermathecal contractions in Locusta migratoria[J].Journal of Insect Physiology,2001,47(6):607-616.
[3] KUBRAKIEWICZ J,JEDRZEJOWKA I,SZYMA?S K A B,et al.Micropyle in neuropterid insects.Structure and late stages of morphogenesis[J].Arthropod Structure & Development,2005,34(2):179-188.
[4] YAMAUCHI H,YOSHITAKE N.Formation and ultrastructure of the micropylar apparatus in Bombyx mori ovarian follicles[J].Journal of Morphology,1984,179(1):47-58.
[5] 夏邦颖.试论昆虫卵的分类特征[J].昆虫分类学报,1980,2(4):247-256.XIA B Y.Taxonomic features of the insect egg[J].Entomotaxonomia,1980,2(4):247-256.
[6] 夏邦颖.昆虫的卵壳[J].生物学通报,2000,35(1):1-3.XIA B Y.Insect shells[J].Bulletin of Biology,2000,35(1):1-3.
[7] ZARANI F E,MARGARITIS L H.The eggshell of the almond wasp Eurytoma amygdale(Hymenoptera,Eurytomidae)-2.The micropylar appendage[J].Tissue & Cell,1994,26(4):569-577.
[8] GULLAN P,CRANSTON P.The Insects:An Outline of Entomology[M].Oxford:Blackwell Publishing,2005.
[9] OSAWA N,YOSHINAGA A.The presence of micropyles in the shells of developing and undeveloped eggs of the ladybird beetle Harmonia axyridis(Coleoptera:Coccinellidae)[J].European Journal of Entomology,2009,106(4):607-610.
[10] REZENDE G L,VARGAS H C M,MOUSSIAN B,et al.Composite eggshell matrices:chorionic layers and sub-chorionic cuticular envelopes[M]// Extracellular Composite Matrices in Arthropods.New York:Springer,2016:325-366.
[11] 彭晨星.家蚕淡墨突变体(bd)雌性不育的分子机制研究[D].重庆:西南大学,2017.PENG CH X.A study on the female infertility of black dilute mutant of silkworm Bombyx mori[D].Chongqing:Southwest University,2017.
[12] ELIANA P L,ICHHA K,PEDRO G,et al.The hippo pathway polarizes the actin cytoskeleton during collective migration of drosophila border cells[J].Journal of Cell Biology,2013,201(6):875-885.
[13] ANDREW R J,TEMBHARE D B.Surface ultrastructure of the egg chorion in the dragonfly,Ictinogomphus rapax (Rambur) (Odonata,Gomphidae)[J].International Journal of Insect Morphology & Embryology,1992,21(4):347-350.
[14] MOUZAKI D G,ZARANI F E,MARGARITIS L H.Structure and morphologenesis of the eggshell and mictopylar apparatus in the olive fly,Dacus oleae(Diptera:Tephritidae)[J].Journal of Morphology,1991,209(1):39-52.
[15] SARASHINA I,MITO T,SAITO M,et al.Location of micropyles and early embryonic development of the two-spotted cricket Gryllus bimaculatus (Insecta,Orthoptera)[J].Development Growth & Differentiation,2005,47(2):99-108.
[16] HINTON H E.Biology of Insect Eggs[M].Oxford:Pergamon Press,1981:1125.
[17] NEDVéD O,HONêK A.Life history and development[M]// HODEK I,VAN EMDEN H F,HON?K A.Ecology of Coccinellidae.Oxford:Wiley-Blackwell,2012:55-56.
[18] MARGARITIS L H.Structure and physiology of the eggshell[M]// GILBERT L I,KERKUT G A.Comprehensive Insect Physiology,Biochemistry and Pharmacology.Vol.1.Oxford:Pergamon Press,1985:153-230.
[19] GAUTAM S G,OPIT G P,MARGOSAN D,et al.Comparative egg morphology and chorionic ultrastructure of key stored-product insect pests[J].Annals of the Entomological Society of America,2015,108(1):43-56.
[20] KAWAGUCHI Y,KUSAKABE T,KOGA K.Morphological variation of micropylar apparatus in Bombyx mori eggs[J].Journal of Insect Biotechnology and Sericology,2002,71(1):49-54.
[21] KUMAR V,KAMBLE C K.Scanning electron microscope study on the egg chorion of silkmoth,Antheraea assamensis Helf.(Lepidoptera:Saturniidae)[J].Animal Biology,2008,58(2):235-244.
[22] ROCHA T,DAVID J A O,HENRIQUE F.Ultramorphological features of the egg of Telmatoscopus albipunctatus(Williston) (Diptera,Psychodidae)[J].Revista Brasileira De Entomologia,2011,55(2):179-182.
[23] LU W,SOUPHANYA P,MONTGOMERY M E.Descriptions of immature stages of Scymnus (Neopullus) sinuanodulus Yu and Yao (Coleoptera:Coccinellidae) with notes on life history[J].Coleopterists Bulletin,2002,56(1):127-141.
[24] YASHIRO T,MATSUURA K.Termite queens close the sperm gates of eggs to switch from sexual to asexual reproduction[J].Proceedings of the National Academy of Sciences of the United States of America,2014,111(48):17212-17217.
[25] MA P W K,BAIRD S,RAMASWAMY S B.Morphology and formation of the eggshell in the tarnished plant bug,Lygus lineloaris (Palisot de Beauvois) (Hemiptera:Miridae)[J].Arthropod Structure & Development,2002,31(2):131-146.
[26] ARTHUR B I,SBILORDO S H,PEMBERTON A J,et al.The anatomy of fertilization in the yellow dung fly Scathophaga stercoraria[J].Journal of Morphology,2008,269(5):630-637.
[27] LEOPOLD R A,DEGRUGILLIER M E.Sperm penetration of housefly eggs:evidence for involvement of female accessory secretion[J].Science,1973,181(4099):555-557.
[28] 夏邦颖.研究昆虫卵壳的实践意义[J].昆虫知识,1982,19(1):25-28.XIA B Y.Practical significance of the study of insect shells[J].Entomological Knowledge,1982,19(1):25-28.
[29] 夏邦颖.几种蛾类卵的受精孔管[J].昆虫分类学报,1984,4(4):325-326.XIA B Y.Micropylar tubes of moth eggs[J].Entomotaxonomia,1984,4(4):325-326.
[30] 夏邦颖.马尾松毛虫卵壳的研究[J].电子显微学报,1984,3(2):58-66.XIA B Y.Studies on the chorion of Dendrolimus punctatus[J].Journal of Chinese Electron Microscopy Society,1984,3(2):58-66.
[31] 花保祯.桃蛀果蛾学名的更正[J].昆虫分类学报,1992,14(4):313-314.HUA B ZH.On the scientific name of the peach fruit borer[J].Entomotaxonomia,1992,14(4):313-314.
[32] 夏邦颖.桃小食心虫卵和稻绿蝽卵的扫描电镜观察[J].昆虫学报,1981,24(1):111-112.XIA B Y.Observation on the structure of egg shells of Carposina niponensis Walsingham and Nezara viridula L.by scanning electron microscopy[J].Acta Entomologica Sinica,1981,24(1):111-112.
[33] CHAPMAN R F.The Insects:Structure and Function [M].Fourth ed.Cambridge:Cambridge University Press,1998.
[34] SANTANA D C,BERGO E S,SALLUM M A M.Anopheles goeldii Rozeboom & Gabaldón (Diptera,Culicidae):a species of the Nuneztovari complex of Anopheles Meigen[J].Revista Brasileira de Entomologia,2015,59:68-76.
[35] KUEROVá Z,STEJSKAL V.External egg morphology of two stored product anobiids,Stegobium paniceum and Lasioderma serricorne(Coleoptera:Anobiidae)[J].Journal of Stored Products Research,2010,46(3):202-205.
[36] KU?EROVá Z,STEJSKAL V.Differences in egg morphology of the stored-grain pests Rhyzopertha dominica and Prostephanus truncatus(Coleoptera:Bostrichidae)[J].Journal of Stored Products Research,2008,44(1):103-105.
[37] VOGELGESANG M,SZKLARZEWICZ T.Formation and structure of egg capsules in scale insects (Hemiptera,Coccinea):I.Ortheziidae[J].Arthropod Structure & Development,2001,30(1):63-68.
[38] 储西平.食蚜蝇科卵壳超微形态初探[J].昆虫学报,1992,35(3):294-300.CHU X P.Preliminary study on the chorionic micromorphology of Syrphidae (Diptera)[J].Acta Entomologica Sinica,1992,35(3):294-300.
[39] KOSS R W.Morphology and taxonomic use of Ephemeroptera eggs[J].Annals of the Entomological Society of America,1968,61(3):696-721.
[40] CHARLAT S,REUTER M,DYSON E A,et al.Male-killing bacteria trigger a cycle of increasing male fatigue and female promiscuity[J].Current Biology,2007,17(3):273-277.
[41] GOWATY P A,KIM Y K,RAWLINGS J,et al.Polyandry increases offspring viability and mother productivity but does not decrease mother survival in Drosophila pseudoobscura[J].Proceedings of the National Academy of Sciences of the United States of America,2010,107(31):13771-13776.
[42] IOSSA G,GAGE M J,EADY P E.Micropyle number is associated with elevated female promiscuity in Lepidoptera[J].Biology Letters,2016,12(12):20160782.
[43] GOBIN B,PEETERS C,BILLEN J.Production of trophic eggs by virgin workers in the ponerine ant Gnamptogenys menadensis[J].Physiological Entomology,1998,23(4):329-336.
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