小菜蛾成虫下唇须感器的超微结构及其神经元中枢投射
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摘要
【目的】明确小菜蛾Plutella xylostella成虫下唇须感器的形态结构及感器神经元的投射。【方法】利用光学显微镜观察和扫描电子显微镜观察下唇须结构和感器类型,利用神经回填技术和激光共聚焦显微镜观察下唇须感器神经元在脑部的投射。【结果】小菜蛾成虫下唇须共3节,其上存在Böhm氏鬃毛、钟形感器、鳞形感器、锥形感器、微毛形感器5种不同类型的感器和一个陷窝器结构。Böhm氏鬃毛短小尖细,钟形感器形如顶部凹陷的圆帽,两种感器均分布于下唇须第1节,且大小上均无雌雄二型差异;鳞形感器形同柳叶,锥形感器粗而直,均散生于下唇须的第2和3节,两种感器在大小上均存在雌雄二型差异,其中雌性的鳞形感器显著大于雄性的,根据其雌雄二型差异现象推测雌蛾的鳞形感器可能与感受寄主植物挥发物有关;下唇须第3节中上部具有一个圆形陷窝器结构,雄虫的陷窝器内径为5.68±0.33μm,雌虫的为6.03±0.23μm,雌雄间无显著性差异;凹坑内长有表面光滑的微毛形感器。小菜蛾下唇须感器神经元主要投射于脑部咽下神经节、每个触角叶的下唇须陷窝器神经纤维球和腹神经索3条通路。【结论】阐明了小菜蛾下唇须感器的类型、分布和形态特征及其感器神经元在脑部的投射形态,为深入了解小菜蛾下唇须感器的生理和功能奠定了基础。
【Aim】 This study aims to characterize the morphology of sensilla on labial palps and the central projection of their sensory neurons in adults of the diamondback moth, Plutella xylostella. 【Methods】 The structures of labial palps and their sensilla type were observed under light and scanning electron microscopy, and the central projection of the sensory neurons of sensilla on labial palps were observed by using neuronal backfills and confocal microscope. 【Results】 The labial palp of P. xylostella adults consists of three segments, on which there are five types of sensilla, including Böhm bristles, sensilla campaniformia, sensilla squamiformia, sensilla basiconica, and microtrichia, and a labial-palp pit organ(LPO). Böhm bristles are spine-like, sensilla campaniformia have circular dome surrounded by a cuticular rim, and both of the two sensilla were found only on the 1 st segment of labial palp, without significant differences in size between sexes. Sensilla squamiformia are willow leaf-like, sensilla basiconica are thick and straight, and both are scattered on the 2 nd and 3 rd segments, with significant differences in size between sexes. Sensilla squamiformia in females are significantly larger than those in males, and this sexual dimorphism suggests that this type of sensilla of female are likely associated with sensing of host plant odours. The LPO is located in the middle and upper part of the 3 rd palpus segment, and the width of the pit is 5.68±0.33 μm in males and 6.03±0.23 μm in females, without significant difference between sexes. Many microtrichia with smooth surface are located in inner pit. The sensory neurons on the labial palp project to the subesophageal ganglion, labial pit organ glomerulus(LPOG) in each antennal lobe and the ventral nerve cord. 【Conclusion】 The morphology of sensilla on labial palps and the central projection of their sensory neurons in P. xylostella adults have been illustrated in this study, providing a basis for further investigation on their physiology and function.
【Aim】 This study aims to characterize the morphology of sensilla on labial palps and the central projection of their sensory neurons in adults of the diamondback moth, Plutella xylostella. 【Methods】 The structures of labial palps and their sensilla type were observed under light and scanning electron microscopy, and the central projection of the sensory neurons of sensilla on labial palps were observed by using neuronal backfills and confocal microscope. 【Results】 The labial palp of P. xylostella adults consists of three segments, on which there are five types of sensilla, including Böhm bristles, sensilla campaniformia, sensilla squamiformia, sensilla basiconica, and microtrichia, and a labial-palp pit organ(LPO). Böhm bristles are spine-like, sensilla campaniformia have circular dome surrounded by a cuticular rim, and both of the two sensilla were found only on the 1 st segment of labial palp, without significant differences in size between sexes. Sensilla squamiformia are willow leaf-like, sensilla basiconica are thick and straight, and both are scattered on the 2 nd and 3 rd segments, with significant differences in size between sexes. Sensilla squamiformia in females are significantly larger than those in males, and this sexual dimorphism suggests that this type of sensilla of female are likely associated with sensing of host plant odours. The LPO is located in the middle and upper part of the 3 rd palpus segment, and the width of the pit is 5.68±0.33 μm in males and 6.03±0.23 μm in females, without significant difference between sexes. Many microtrichia with smooth surface are located in inner pit. The sensory neurons on the labial palp project to the subesophageal ganglion, labial pit organ glomerulus(LPOG) in each antennal lobe and the ventral nerve cord. 【Conclusion】 The morphology of sensilla on labial palps and the central projection of their sensory neurons in P. xylostella adults have been illustrated in this study, providing a basis for further investigation on their physiology and function.
引文
Badenes-Perez FR., Reichelt M, Gershenzon J, Heckel DG, 2013. Interaction of glucosinolate content of Arabidopsis thaliana, mutant lines and feeding and oviposition by generalist and specialist lepidopterans. Phytochemistry, 86(2): 36-43.
Barcaba T, Krenn HW, 2015. The mouthparts of adult Indian meal moths, Plodia interpunctella (Hübner, 1813) (Lepidoptera: Pyralidae). Entomol. Aust., 22: 91-105.
Bogner F, Boppré M, Ernst KD, Boeckh J, 1986. CO2 sensitive receptors on labial palps of Rhodogastria moths (Lepidoptera: Arctiidae): physiology, fine structure and central projection. J. Comp. Physiol. A, 158(6): 741-749.
Chen J, Hua B, 2016. Sexual dimorphism of adult labial palps of the peach fruit moth Carposina sasakii Matsumura (Lepidoptera: Carposinidae) with notes on their sensilla. Acta Zool., 97(1): 42-48.
Couto A, Alenius M, Dickson BJ, 2005. Molecular, anatomical, and functional organization of the Drosophila olfactory system. Curr. Biol., 15: 1535-1547.
Dong JF, Liu H, Tang QB, Liu Y, Zhao XC, Wang GR, 2014. Morphology, type and distribution of the labial-palp pit organ and its sensilla in the oriental armyworm, Mythimna separata (Lepidoptera: Noctuidae). Acta Entomol. Sin., 57(6): 681-687. [董钧锋, 刘辉, 汤清波, 刘杨, 赵新成, 王桂荣, 2014. 粘虫下唇须陷窝器及其感器的形态、类型与分布. 昆虫学报, 57(6): 681-687]
Faucheux MJ, 2005. Heterobathmia pseuderiocrania (Heterobathmiina): mouthparts, mouthpart sensilla, and comparison with other non-glossatan suborders (Insecta, Lepidoptera). Rev. Mus. Argent. Cienc. Nat., n. s., 7(1): 57-65.
Faucheux MJ, 2008. Mouthparts and associated sensilla of a South American moth, Synempora andesae (Lepidoptera: Neopseustidae). Rev. Soc. Entomol. Argent., 67(1-2): 21-33.
Faucheux MJ, 2012. Wall-pore sensilla auricillica on the labial palps of Limnephilus marmoratus Curtis 1834 (Trichoptera: Limnephilidae). Ann. Soc. Entomol. Fr. (n.s.), 48 (1-2): 229-231.
Galizia CG, R?ssler W, 2010. Parallel olfactory systems in insects: anatomy and function. Annu. Rev. Entomol., 55: 399-420.
Guerenstein PG, Christensen TA, Hildebrand JG, 2004a. Sensory processing of ambient CO2 information in the brain of the moth Manduca sexta. J. Comp. Physiol. A, 190: 707-725.
Guerenstein PG, Yepez EA, van Haren J, Williams DG, Hildebrand JG, 2004b. Floral CO2 emission may indicate food abundance to nectar-feeding moths. Naturwissenschaften, 91(7): 329-333.
Huang J, Ma JW, Liu XR, Li MM, Hua BZ, 2011. Ultrastructure of the mouthpart sensilla in adult Bittacus sinensis Walker (Mecoptera: Bittacidae). Acta Entomol. Sin., 54(1): 110-116. [黄晶, 马继文, 刘歆然, 李美美, 花保祯, 2011. 中华蚊蝎蛉成虫口器感器的超微结构. 昆虫学报, 54(1): 110-116]
Kent KS, Harrow ID, Quartararo P, Hildebrand JG, 1986. An accessory olfactory pathway in Lepidoptera: the labial pit organ and its central projections in Manduca sexta and certain other sphinx moths and silk moths. Cell Tissue Res., 245(2): 237-245.
Krenn HW, Penz CM, 1998. Mouthparts of Heliconius butterflies (Lepidoptera: Nymphalidae): a search for anatomical adaptations to pollen-feeding behavior. Int. J. Insect Morphol. Embryol., 27(4): 301-309.
Lee JK, Altner H, 1986. Primary sensory projections of the labial palp-pit organ of Pieris rapae L. (Lepidoptera: Pieridae). Int. J. Insect Morphol. Embryol., 15(5-6): 439-448.
Lee JK, Selzer R, Altner H, 1985. Lamellated outer dendritic segments of a chemoreceptor within wall-pore sensilla in the labial palp-pit organ of the butterfly, Pieris rapae L. (Insecta, Lepidoptera). Cell Tissue Res., 240(2): 333-342.
Malo EA, Castrejón-Gómez VR, Cruz-López L, Rojas JC, 2004. Antennal sensilla and electrophysiological response of male and female Spodoptera frugiperda (Lepidoptera: Noctuidae) to conspecific sex pheromone and plant odors. Ann. Entomol. Soc. Am., 97(6): 1273-1284.
McIver SB, 1975. Structure of cuticular mechanoreceptors of arthropods. Annu. Rev. Entomol., 20: 381-397.
Merivee E, Ploomi A, Rahi M, Bresciani J, Ravn HP, Luik A, Sammelselge V, 2002. Antennal sensilla of the ground beetle Bembidion properans Steph. (Coleoptera, Carabidae). Micron, 33(5): 429-440.
Ning C, Yang K, Xu M, Huang LQ, Wang CZ, 2016. Functional validation of the carbon dioxide receptor in labial palps of Helicoverpa armigera, moths. Insect Biochem. Molec. Biol., 73: 12-19.
Schneider D, 1964. Insect antennae. Annu. Rev. Entomol., 9: 103-122.
Shields VD, Hildebrand JG, 2001. Recent advances in insect olfaction, specifically regarding the morphology and sensory physiology of antennal sensilla of the female sphinx moth Manduca sexta. Microsc. Res. Tech., 55(5): 307-329.
Song YQ, Sun HZ, Wu JX, 2016. Ultrastructural characteristics of the proboscis and the labial palp pit organ in the oriental fruit moth, Grapholita molesta. Bull. Insectol., 69(1): 59-66.
Stange G, 1997. Effects of changes in atmospheric carbon dioxide on the location of hosts by the moth, Cactoblastis cactorum. Oecologia, 110: 539-545.
Stange G, Monro J, Stowe S, Osmond CB, 1995. The CO2 sense of the moth Cactoblastis cactorum, and its probable role in the biological control of the CAM plant Opuntia stricta. Oecologia, 102: 341-352.
Sun JY, S?nderby IE, Halkier BA, Jander G, de Vos M, 2009. Non-volatile intact indole glucosinolates are host recognition cues for ovipositing Plutella xylostella. J. Chem. Ecol., 35(12): 1427-1436.
Sun X, Wang MQ, Zhang G, 2011. Ultrastructural observations on antennal sensilla of Cnaphalocrocis medinalis (Lepidoptera: Pyralidae). Microsc. Res. Tech., 74(2): 113-121.
Thom C, Guerenstein PG, 2004. Floral CO2 reveals flower profitability to moths. J. Chem. Ecol., 30(6): 1285-1288.
Varela N, Couton L, Gemeno C, Avilla J, Rospars JP, Anton S, 2009. Three-dimensional antennal lobe atlas of the oriental fruit moth, Cydia molesta (Busck) (Lepidoptera: Tortricidae): comparison of male and female glomerular organization. Cell Tissue Res., 337: 513-526.
Wee SL, Oh HW, Park KC, 2016. Antennal sensillum morphology and electrophysiological responses of olfactory receptor neurons in trichoid sensilla of the diamondback moth (Lepidoptera: Plutellidae). Fla. Entomol., 99: 146-158.
Yan XZ, Deng CP, Sun XJ, Hao C, 2014. Effects of various degrees of antennal ablation on mating and oviposition preferences of the diamondback moth, Plutella xylostella L. J. Integr. Agr., 13(6):1311-1319.
Yan XZ, Deng CP, Xie JX, Wu LJ, Sun XJ, Hao C, 2017. Distribution patterns and morphology of sensilla on the antennae of Plutella xylostella (L.) -a scanning and transmission electron microscopic study. Micron, 103: 1-11.
Yan XZ, Xie JX, Deng CP, Wu LJ, Sun XJ, Hao C, 2017. Number, distribution and morphology of cell bodies in the antennal lobe of Plutella xylostella (Lepidoptera: Plutellidae) adults. Acta Entomol. Sin., 60(7): 735-741. [闫喜中, 谢佼昕, 邓彩萍, 吴兰军, 孙学俊, 郝赤, 2017. 小菜蛾成虫触角叶细胞体的数量、分布和形态. 昆虫学报, 60(7): 735-741]
Yang H, Yan SC, Liu D, 2009. Ultrastructural observations on antennal sensilla of Coleophora obducta (Meyrick) (Lepidoptera: Coleophoridae). Micron, 40: 231-238.
Zhao XC, Tang QB, Berg BG, Liu Y, Wang YR, Yan FM, Wang GR, 2013. Fine structure and primary sensory projections of sensilla located in the labial-palp pit organ of Helicoverpa armigera (Insecta). Cell Tissue Res., 353: 399-408.
Barcaba T, Krenn HW, 2015. The mouthparts of adult Indian meal moths, Plodia interpunctella (Hübner, 1813) (Lepidoptera: Pyralidae). Entomol. Aust., 22: 91-105.
Bogner F, Boppré M, Ernst KD, Boeckh J, 1986. CO2 sensitive receptors on labial palps of Rhodogastria moths (Lepidoptera: Arctiidae): physiology, fine structure and central projection. J. Comp. Physiol. A, 158(6): 741-749.
Chen J, Hua B, 2016. Sexual dimorphism of adult labial palps of the peach fruit moth Carposina sasakii Matsumura (Lepidoptera: Carposinidae) with notes on their sensilla. Acta Zool., 97(1): 42-48.
Couto A, Alenius M, Dickson BJ, 2005. Molecular, anatomical, and functional organization of the Drosophila olfactory system. Curr. Biol., 15: 1535-1547.
Dong JF, Liu H, Tang QB, Liu Y, Zhao XC, Wang GR, 2014. Morphology, type and distribution of the labial-palp pit organ and its sensilla in the oriental armyworm, Mythimna separata (Lepidoptera: Noctuidae). Acta Entomol. Sin., 57(6): 681-687. [董钧锋, 刘辉, 汤清波, 刘杨, 赵新成, 王桂荣, 2014. 粘虫下唇须陷窝器及其感器的形态、类型与分布. 昆虫学报, 57(6): 681-687]
Faucheux MJ, 2005. Heterobathmia pseuderiocrania (Heterobathmiina): mouthparts, mouthpart sensilla, and comparison with other non-glossatan suborders (Insecta, Lepidoptera). Rev. Mus. Argent. Cienc. Nat., n. s., 7(1): 57-65.
Faucheux MJ, 2008. Mouthparts and associated sensilla of a South American moth, Synempora andesae (Lepidoptera: Neopseustidae). Rev. Soc. Entomol. Argent., 67(1-2): 21-33.
Faucheux MJ, 2012. Wall-pore sensilla auricillica on the labial palps of Limnephilus marmoratus Curtis 1834 (Trichoptera: Limnephilidae). Ann. Soc. Entomol. Fr. (n.s.), 48 (1-2): 229-231.
Galizia CG, R?ssler W, 2010. Parallel olfactory systems in insects: anatomy and function. Annu. Rev. Entomol., 55: 399-420.
Guerenstein PG, Christensen TA, Hildebrand JG, 2004a. Sensory processing of ambient CO2 information in the brain of the moth Manduca sexta. J. Comp. Physiol. A, 190: 707-725.
Guerenstein PG, Yepez EA, van Haren J, Williams DG, Hildebrand JG, 2004b. Floral CO2 emission may indicate food abundance to nectar-feeding moths. Naturwissenschaften, 91(7): 329-333.
Huang J, Ma JW, Liu XR, Li MM, Hua BZ, 2011. Ultrastructure of the mouthpart sensilla in adult Bittacus sinensis Walker (Mecoptera: Bittacidae). Acta Entomol. Sin., 54(1): 110-116. [黄晶, 马继文, 刘歆然, 李美美, 花保祯, 2011. 中华蚊蝎蛉成虫口器感器的超微结构. 昆虫学报, 54(1): 110-116]
Kent KS, Harrow ID, Quartararo P, Hildebrand JG, 1986. An accessory olfactory pathway in Lepidoptera: the labial pit organ and its central projections in Manduca sexta and certain other sphinx moths and silk moths. Cell Tissue Res., 245(2): 237-245.
Krenn HW, Penz CM, 1998. Mouthparts of Heliconius butterflies (Lepidoptera: Nymphalidae): a search for anatomical adaptations to pollen-feeding behavior. Int. J. Insect Morphol. Embryol., 27(4): 301-309.
Lee JK, Altner H, 1986. Primary sensory projections of the labial palp-pit organ of Pieris rapae L. (Lepidoptera: Pieridae). Int. J. Insect Morphol. Embryol., 15(5-6): 439-448.
Lee JK, Selzer R, Altner H, 1985. Lamellated outer dendritic segments of a chemoreceptor within wall-pore sensilla in the labial palp-pit organ of the butterfly, Pieris rapae L. (Insecta, Lepidoptera). Cell Tissue Res., 240(2): 333-342.
Malo EA, Castrejón-Gómez VR, Cruz-López L, Rojas JC, 2004. Antennal sensilla and electrophysiological response of male and female Spodoptera frugiperda (Lepidoptera: Noctuidae) to conspecific sex pheromone and plant odors. Ann. Entomol. Soc. Am., 97(6): 1273-1284.
McIver SB, 1975. Structure of cuticular mechanoreceptors of arthropods. Annu. Rev. Entomol., 20: 381-397.
Merivee E, Ploomi A, Rahi M, Bresciani J, Ravn HP, Luik A, Sammelselge V, 2002. Antennal sensilla of the ground beetle Bembidion properans Steph. (Coleoptera, Carabidae). Micron, 33(5): 429-440.
Ning C, Yang K, Xu M, Huang LQ, Wang CZ, 2016. Functional validation of the carbon dioxide receptor in labial palps of Helicoverpa armigera, moths. Insect Biochem. Molec. Biol., 73: 12-19.
Schneider D, 1964. Insect antennae. Annu. Rev. Entomol., 9: 103-122.
Shields VD, Hildebrand JG, 2001. Recent advances in insect olfaction, specifically regarding the morphology and sensory physiology of antennal sensilla of the female sphinx moth Manduca sexta. Microsc. Res. Tech., 55(5): 307-329.
Song YQ, Sun HZ, Wu JX, 2016. Ultrastructural characteristics of the proboscis and the labial palp pit organ in the oriental fruit moth, Grapholita molesta. Bull. Insectol., 69(1): 59-66.
Stange G, 1997. Effects of changes in atmospheric carbon dioxide on the location of hosts by the moth, Cactoblastis cactorum. Oecologia, 110: 539-545.
Stange G, Monro J, Stowe S, Osmond CB, 1995. The CO2 sense of the moth Cactoblastis cactorum, and its probable role in the biological control of the CAM plant Opuntia stricta. Oecologia, 102: 341-352.
Sun JY, S?nderby IE, Halkier BA, Jander G, de Vos M, 2009. Non-volatile intact indole glucosinolates are host recognition cues for ovipositing Plutella xylostella. J. Chem. Ecol., 35(12): 1427-1436.
Sun X, Wang MQ, Zhang G, 2011. Ultrastructural observations on antennal sensilla of Cnaphalocrocis medinalis (Lepidoptera: Pyralidae). Microsc. Res. Tech., 74(2): 113-121.
Thom C, Guerenstein PG, 2004. Floral CO2 reveals flower profitability to moths. J. Chem. Ecol., 30(6): 1285-1288.
Varela N, Couton L, Gemeno C, Avilla J, Rospars JP, Anton S, 2009. Three-dimensional antennal lobe atlas of the oriental fruit moth, Cydia molesta (Busck) (Lepidoptera: Tortricidae): comparison of male and female glomerular organization. Cell Tissue Res., 337: 513-526.
Wee SL, Oh HW, Park KC, 2016. Antennal sensillum morphology and electrophysiological responses of olfactory receptor neurons in trichoid sensilla of the diamondback moth (Lepidoptera: Plutellidae). Fla. Entomol., 99: 146-158.
Yan XZ, Deng CP, Sun XJ, Hao C, 2014. Effects of various degrees of antennal ablation on mating and oviposition preferences of the diamondback moth, Plutella xylostella L. J. Integr. Agr., 13(6):1311-1319.
Yan XZ, Deng CP, Xie JX, Wu LJ, Sun XJ, Hao C, 2017. Distribution patterns and morphology of sensilla on the antennae of Plutella xylostella (L.) -a scanning and transmission electron microscopic study. Micron, 103: 1-11.
Yan XZ, Xie JX, Deng CP, Wu LJ, Sun XJ, Hao C, 2017. Number, distribution and morphology of cell bodies in the antennal lobe of Plutella xylostella (Lepidoptera: Plutellidae) adults. Acta Entomol. Sin., 60(7): 735-741. [闫喜中, 谢佼昕, 邓彩萍, 吴兰军, 孙学俊, 郝赤, 2017. 小菜蛾成虫触角叶细胞体的数量、分布和形态. 昆虫学报, 60(7): 735-741]
Yang H, Yan SC, Liu D, 2009. Ultrastructural observations on antennal sensilla of Coleophora obducta (Meyrick) (Lepidoptera: Coleophoridae). Micron, 40: 231-238.
Zhao XC, Tang QB, Berg BG, Liu Y, Wang YR, Yan FM, Wang GR, 2013. Fine structure and primary sensory projections of sensilla located in the labial-palp pit organ of Helicoverpa armigera (Insecta). Cell Tissue Res., 353: 399-408.