草蛉滞育的研究进展

doi:10.16409/j.cnki.2095-039x.2019.03.006

摘要/Abstract

摘要： 草蛉是多种农林害虫的重要捕食性天敌昆虫，在生物防治中具有重要作用。滞育是昆虫应对不利环境条件时，生长发育停滞的一种生理状态，许多草蛉种类存在滞育现象。深入开展草蛉滞育的研究，不仅有助于明确其滞育特征和内在调控机制，也有助于提高草蛉的生物防治效果，促进草蛉的商业化生产和应用。本文归纳总结了1910年以来国内外草蛉科种类的滞育研究文献，介绍了目前已开展滞育研究的草蛉种类及其滞育特征，分析了影响草蛉滞育调控的环境因子，论述了草蛉滞育期间的形态和生理生化变化、分子机制及其滞育后发育生物学，并就滞育在草蛉规模化生产中的应用以及目前草蛉滞育研究中存在的问题进行了讨论，以期为深入草蛉滞育的基础理论研究和促进草蛉的规模化生产应用等提供参考依据。

关键词: 草蛉, 滞育, 环境因子, 生理变化, 滞育后发育生物学

Abstract: The green lacewings are important predatory natural enemies of a variety of small, soft-bodied pests in agro-ecological systems and forests. Many species of green lacewings enter diapause in response to adverse environmental conditions. Diapause research of the green lacewings is essential for understanding characteristics and regulation mechanisms of diapause and for enhancing the biocontrol potential, thus promoting the commercial production and application of green lacewings. In this review, we summarized the literatures of diapause in the chrysopids since 1910, which included the diapause characteristics, environmental factors responsible for diapause regulation, the behavioral and physiological changes during diapause, the molecular mechanisms of diapause, and the biology of post-diapause development. The application of diapause in the mass rearing of chrysopids and research prospectives were discussed. This review serves to further the studies of diapause of chrysopids and to facilitate large-scale production and application of chrysopids in biological control.

Key words: chrysopids, diapause, environmental factors, physiological changes, post-diapause development

中图分类号:

S476.2

王曼姿, 李玉艳, 高飞, 张礼生. 草蛉滞育的研究进展[J]. 中国生物防治学报, 2019, 35(3): 474-486.

WANG Manzi, LI Yuyan, GAO Fei, ZHANG Lisheng. Research Advances in Diapause of Green Lacewings (Neuroptera: Chrysopidae)[J]. journal1, 2019, 35(3): 474-486.

参考文献

[1] Tauber M J, Tauber C A, Masaki S. Seasonal Adaptations of Insects[M]. Oxford:Oxford University Press, 1986.
[2] Danks H V. Insect Dormancy:An Ecological Perspective[M]. Ottawa:Biological Survey of Canada (Terrestrial Arthropods), 1987, 439.
[3] Kostál V. Eco-physiological phases of insect diapause[J]. Journal of Insect Physiology, 2006, 52(2):113-127.
[4] Denlinger D L, Yocum G D, Rinehart J P. 10-Hormonal Control of Diapause[M]//Gilbert L I ed. Insect Endocrinology. Elsevier/Academic Press, 2012, 430-463.
[5] 张礼生, 曾凡荣. 滞育和休眠在昆虫饲养中的应用[M]//曾凡荣, 陈红印, 主编. 天敌昆虫饲养系统工程. 北京:中国农业科学技术出版社, 2009, 54-89.
[6] 李玉艳, 张礼生, 陈红印. 茧蜂滞育的研究进展[J]. 昆虫学报, 2010, 53(10):1167-1178.
[7] McEwen P K, New T R, Whittington A E. Lacewings in the crop environment[J]. Zoological Journal of the Linnean Society, 2010, 151(3):661-661.
[8] 杨集昆. 草蛉的生活习性和常见种类[J]. 应用昆虫学报, 1974, 11(3):38-43.
[9] 杨星科, 杨集昆, 李文柱. 中国动物志:昆虫纲, 第三十九卷, 脉翅目, 草蛉科[M]. 北京:科学出版社, 2005, 1-398.
[10] 武鸿鹄. 温室环境因子对大草蛉和丽草蛉成虫扩散行为的影响研究[D]. 北京:中国农业科学院, 2014.
[11] 蒋佩兰, 朱杏芬, 章志英. 南昌地区主要草蛉的习性及优势种利用研究初探[J]. 江西农业大学学报, 1993, 15(4):467-470.
[12] Cranshaw W, Sclar D C, Cooper D. A review of 1994 pricing and marketing by suppliers of organisms for biological control of arthropods in the United States[J]. Biological Control Theory and Applications in Pest Management, 1996, 6(2):291-296.
[13] Lenteren J C V, Roskam M M, Timmer R. Commercial mass production and pricing of organisms for biological control of pests in Europe[J]. Biological Control, 1997, 10(2):143-149.
[14] Yang N W, Zang L S, Wang S, et al. Biological pest management by predators and parasitoids in the greenhouse vegetables in China[J]. Biological Control, 2014, 68(1):92-102.
[15] 张礼生, 陈红印. 生物防治作用物研发与应用的进展[J]. 中国生物防治学报, 2014, 30(5):581-586.
[16] 张帆, 李姝, 肖达, 等. 中国设施蔬菜害虫天敌昆虫应用研究进展[J]. 中国农业科学, 2015, 48(17):3463-3476.
[17] 邵振芳, 尹文兵, 陈建华, 等. 草蛉在虫害生物防治中的应用研究进展[J]. 现代农业科技, 2016(3):171-174.
[18] New T R. The biology of Chrysopidae and Hemerobiidae (Neuroptera), with reference to their usage as biocontrol agents:a review[J]. Ecological Entomology, 1975, 127(2):115-140.
[19] Canard M, Principi M M. Development of chrysopidae[M]//Canard M, Séméria Y, New T R, eds. Biology of Chrysopidae. The Hague:Dr. W. Junk, 1984, 294.
[20] Nordulund D A, Correa J A. Improvements in the production system for green lacewings:an adult feeding and oviposition unit and hot wire egg harvesting system[J]. Biological Control, 1995, 5(2):179-188.
[21] 张帆, 王素琴, 罗晨, 等. 几种人工饲料及繁殖技术对大草蛉生长发育的影响[J]. 植物保护, 2004, 30(5):36-40.
[22] 林美珍, 陈红印, 王树英, 等. 大草蛉幼虫人工饲料的研究[J]. 中国生物防治学报, 2007, 23(4):316-321.
[23] 陈珍珍, 李明贵, 郭亚楠, 等. 光周期和温度对中华通草蛉不同越冬时期成虫滞育后生物学特性的影响[J]. 中国农业科学, 2013, 46(8):1610-1618.
[24] Canard M. Seasonal adaptations of green lacewings (Neuroptera:Chrysopidae)[J]. European Journal of Entomology, 2005, 102(3):317-324.
[25] Volkovich T A. Seasonal development of a lacewing, Chrysopa dorsalis Burmeister (Neuroptera:Chrysopidae) in the forest-steppe zone of Russia[J]. Entomologicheskoe Obozrenie, 2006, 85(3):473-486.
[26] Li Y Y, Wang M Z, Gao F, et al. Exploiting diapause and cold tolerance to enhance the use of the green lacewing Chrysopa formosa Brauer in biological control[J]. Biological Control, 2018, 127(1):116-126.
[27] Volkovich T A. Environmental control of seasonal cycles in green lacewings (Neuroptera, Chrysopidae) from the forest-Steppe Zone of Russia[J]. Acta Zoologica Fennica, 1998, 209:263-275.
[28] Tauber M J, Tauber C A. Larval diapause in Chrysopa nigricornis:sensitive stages, critical photoperiod, and termination (Neuroptera:Chrysopidae)[J]. Entomologia Experimentalis et Applicata, 1972, 15(1):105-111.
[29] Nakahira K, Arakawa R. Effect of photoperiod on the development and diapause of the green lacewing Chrysopa pallens (Neuroptera:Chrysopidae)[J]. Entomological Science, 2005, 8(2):133-135.
[30] Kharizanov A, Babrikova T. A study of some features of the biology of Chrysopa septempunctata[J]. Rastitelna Zashchita, 1976, 24(10):26-30.
[31] Canard M. Diapause in Chrysopa perla (L) (Neuroptera Chrysopidae) induction and elimination in natural and experimental conditions[J]. Annales de Zoologie Ecologie Animale, 1976, 8(1):10-19.
[32] Zelený J. Lace-wings (Neuroptera) in cultual steppe and the population dynamics in the species Chrysopa carnea Stephens and Chrysopa phyllochroma Wesmael[J]. Acta Entomological Bohemoslovaca, 1965, 62:177-194.
[33] 牟吉元, 王念慈, 范永贵. 四种草蛉生活史和习性的研究[J]. 植物保护学报, 1980, 7(1):1-8.
[34] Sagné J C. Glucidic variations in the lacewing Chrysopa walkeri during the prepupal diapause[J]. Entomologia Experimentalis et Applicata, 1986, 41(1):101-103.
[35] Macleod E G. Experimental induction and elimination of adult diapause and autumnal coloration in Chrysopa carnea (Neuroptera)[J]. Journal of Insect Physiology, 1967, 13(9):1343-1349.
[36] Tauber M J, Tauber C A. Quantitative response to daylength during diapause in insects[J]. Nature, 1973, 244(5414):296-297.
[37] Hodek I. Role of environmental factors and endogenous mechanisms in the seasonality of reproduction in insects diapausing as adults[M]//Brown V K, Hodek I, eds. Diapause and Life Cycle Strategies in Insects. The Huge:Dr.W. Junk, 1983, 9-33.
[38] Tauber M J, Tauber C A. Nutritional and photoperiodic control of the seasonal reproductive cycle in Chrysopa Mohave (Neuroptera)[J]. Journal of Insect Physiology, 1973, 19(4):729-736.
[39] Tauber C A, Tauber M J. Environmental control of univoltinism and its evolution in an insect species[J]. Canadian Journal of Zoology, 1976, 54(2):260-265.
[40] Albuquerque G S, Tauber C A, Tauber M J. Chrysoperla externa (Neuroptera:Chrysopidae):life history and potential for biological control in central and south America[J]. Biological Control, 1994, 4(1):8-13.
[41] Tauber M J, Tauber C A. Developmental requirements of the univoltine species Chrysopa downesi:photoperiodic stimuli and sensitive stages[J]. Journal of Insect Physiology, 1976, 22(2):331-335.
[42] Volkovich T A, Sokolova I V. Thermoperiodic control of diapause in two green lacewing species (Neuroptera:Chrysopidae):a comparative effect of abrupt and gradual changes in temperature[J]. ?ntomologicheskoe Obozrenie, 2000, 79(4):753-761.
[43] Xu Y Y, Mu J, Cui H, et al. Photoperiodic control of adult diapause in chrysoperla sinica (Tjeder) (Neuroptera:Chrysopodae)-I.crirical photoperiod and sensitive stages of adult diapause induction[J]. Insect Science, 2004, 11(3):191-198.
[44] 陈珍珍. 中华通草蛉对长、短光周期的响应研究[D]. 济南:山东农业大学, 2014.
[45] 许永玉, 胡萃, 牟吉元, 等. 中华通草蛉成虫越冬体色变化与滞育的关系[J]. 生态学报, 2002, 22(8):1275-1280.
[46] Chen Z Z, Liu L Y, Liu S Y, et al. Response of Chrysoperla nipponensis (Okamoto) (Neuroptera:Chrysopidae) under long and short photoperiods[J]. Journal of Insect Science, 2017, 17(2):35.
[47] Principi M M, Castellari P L. Larval diapause in Anisochrysa flavifrons (Brauer) (Neuroptera Chrysopidae)[J]. Atti della Accademia delle scienze dell'Istituto di Bologna (Classe di scienze fisiche, 12), 1970, 7:75-83.
[48] Grimal A, Canard M. Preliminary observations on the effect of photoperiod on the life cycle of the green lacewing Hypochrysa elegans (Burmeister) (Insecta:Neuroptera:Chrysopidae:Nothochrysinae)[C]//Canard M, Aspöck H, Mansell M W, eds. Pure and Applied Research in Neuropterology. Proceedings of the Fifth International Symposium on Neuropterology. Cairo, Egypt, 1994. Imprimerie Sacco, Toulouse, France, 1996, 119-127.
[49] Tauber C A, TeresaDeLeón, Lopezarroyo J I, et al. Ceraeochrysa placita (Neuroptera:Chrysopidae):generic characteristics of larvae, larval descriptions, and life cycle[J]. Annals of the Entomological Society of America, 1998, 91(5):608-618.
[50] Canard M, Grimal A. Insect photoperiodism:various ways of regulating univoltinism in lacewings (Planipennia:Chrysopidae)[J]. Experientia, 1988, 44(6):523-525.
[51] Canard M. Photoperiodic sensitivity in larvae of the chrysopid Nineta flava[J]. Entomologia Experimentalis et Applicata, 1983, 34(1):111-118.
[52] Canard M. Effect of photoperiod on the first-instar development in the lacewing Nineta pallida[J]. Physiological Entomology, 2010, 15(2):137-140.
[53] Principi M M, Sgobba D. Larval diapause in Mallada clathratus (Schneider) (Neuroptera, Chrysopidae)[J]. Bollettino dell'Istituto di Entomologia ‘Guido Grandi’ della Universita degli Studi di Bologna, 1994(48):75-91.
[54] Tauber M J, Tauber C A. Phenological responses of Pseudomallada (Neuroptera:Chrysopidae):Comparative data from three Nearctic species and interspecific hybrids[J]. European Journal of Entomology, 2015, 112(1):49-62.
[55] Principi M M, Sgobba D. Larval diapause in Mallada (=Anisochrysa) flavifrons (Brauer) (Neuroptera Chrysopidae):photoperiodic cycles responsible for induction and development of diapause, and reactivation and weight gain of the diapausing instar[J]. Bollettino dell'Istituto di Entomologia ‘Guido Grandi’ della Universita degli Studi di Bologna, 1987(41):209-231.
[56] Canard M. Long rather than short diapause-promoting photoperiods induce strong diapause in green lacewings (Neuroptera:Chrysopidae)[J]. Acta Societatis Zoologicae Bohemicae, 2001, 65(3):163-168.
[57] Denlinger D L. Why study diapause[J]. Entomological Research, 2008, 38(1):1-9.
[58] 时爱菊, 徐洪富, 刘忠德, 等. 光周期对大草蛉(Chrysopa pallens)滞育及发育的影响[J]. 生态学报, 2008, 28(8):3854-3859.
[59] 许永玉. 中华通草蛉的滞育机制和应用研究[D]. 杭州:浙江大学, 2001.
[60] Canard M, Volkovich T A. Outlines of lacewing development[M]//McEwen P, New T R, Whittington A E, eds. Lacewings in the Crop Environment. Cambridge:Cambridge University Press, 2001, 130-153.
[61] 程丽媛. 替代猎物、光周期和温度对大草蛉滞育维持、滞育解除及发育的影响[D]. 济南:山东农业大学, 2017.
[62] Beck S D. Effects of thermoperiod on photoperiodic determination of larval diapause in Ostrinianubilalis[J]. Journal of Insect Physiology, 1985, 31(1):41-46.
[63] 李玉艳, 陈红印, 王孟卿, 等. 环境因子对寄生蜂滞育的影响[C]. 粮食安全与植保科技创新, 2009.
[64] Tauber M J, Tauber C A. Geographic variation in critical photoperiod and in diapause intensity of Chrysopa carnea (Neuroptera)[J]. Journal of Insect Physiology, 1972, 18(1):25-29.
[65] 时爱菊. 大草蛉滞育特性的研究[D]. 济南:山东农业大学, 2007.
[66] 巫国瑞译. 昆虫迁飞和滞育的进化[M]. 北京:科学出版社, 1984.
[67] Orlova N A. Effects of photoperiod and temperature on diapause induction in Chrysopa pallens (Rambur) (Neuroptera:Chrysopidae)[J]. Acta Zoologica Fennica, 1998, 209:195-202.
[68] Canard M, Séméria Y, New T R. Biology of Chrysopidae[M]. Hague:Dr. W. Junk, 1984.
[69] Volkovich T A, Blumental N A. Photo-thermoperiodic responses in some species of lacewings (Neuroptera:Chrysopidae):their role in diapause induction[J]. European Journal of Entomology, 1997, 94(4):435-444.
[70] Masaki S. Summer diapause[J]. Annal Review of Entomology, 1980, 25:1-25.
[71] Tauber M J, Tauber C A. Seasonal responses and their geographic variation in Chrysopa downesi:ecophysiological and evolutionary considerations[J]. Canadian Journal of Zoology, 1981, 59(3):370-376.
[72] Tauber M J, Tauber C A. Inheritance of photoperiodic responses controlling diapause[J]. Bulletin of the Entomological Society of America, 1979, 25(2):125-128.
[73] 杨秋生. 丝带凤蝶滞育与非滞育蛹及其成虫的形态学观察[J]. 昆虫学报, 2008, 51(4):454-458.
[74] Duelli P, Johnson J B, Waldburger M, et al. A new look at adaptive body coloration and color change in "common green lacewings" of the genus Chrysoperla (Neuroptera:Chrysopidae)[J]. Annals of the Entomological Society of America, 2014, 107(2):382-388.
[75] Honek A. Relationship of colour changes and diapause in natural populations of Chrysopa carnea Steph (Neuroptera, Chyrsopidae)[J]. Acta Entomological Bohemoslovaca, 1973, 70(4):254-258.
[76] 许永玉, 牟吉元, 胡萃. 激素对成虫滞育的调控机制[J]. 山东农业大学学报, 1998, 29:17-22.
[77] 李丽英. 昆虫分析生物化学[M]. 北京:科学出版社, 1984, 228-230.
[78] Honek A. Maintenance and termination of winter coloration in Chrysopa carnea (Neuroptera, Chrysopidae):relationships to the development of diapause[J]. Annales de Zoologie Ecologie Animale, 1976, 3(2):114-120.
[79] Denlinger D L. Relationship between cold hardiness and diapause[M]//Lee Jr. R E, Denlinger D L, eds. Insects at Low Temperature. New York:Chapman & Hall, 1991, 174-198.
[80] King A M, Macrae T H. Insect heat shock proteins during stress and diapause[J]. Annual Review of Entomology, 2015, 60(1):59-75.
[81] Principi M M, Memmi M, Sgobba D. Influence of temperature on the larval diapause of Mallada flavifrons (Brauer) (Neuroptera Chrysopidae)[J]. Bollettino dell'Istituto di Entomologia "Guido Grandi", Universita degli Studi Bologna, 1990, 44:37-55.
[82] Canard M, Grimal A, Carvalho C F. Weight changes during preimaginal development in green lacewings (Insecta:Neuroptera:Chrysopidae)[C]//Canard M, Aspöck H, Mansell M W, eds. Pure and Applied Research in Neuropterology. Proceedings of the Fifth International Symposium on Neuropterology. Gairo, Egypt, 1996, 87-101.
[83] 郭海波. 中华通草蛉成虫越冬与滞育的生理生化机制[D]. 济南:山东农业大学, 2006.
[84] 王伟, 张礼生, 陈红印, 等. 瓢虫滞育的研究进展[J]. 植物保护, 2011, 37(5):27-33.
[85] 陈珍珍, 刘力源, 刘韶业, 等. 母代越冬经历对中华通草蛉子代发育与繁殖的影响[J]. 昆虫学报, 2017, 60(5):553-561.
[86] 郅伦山, 李淑芳, 刘兴峰, 等. 中华通草蛉越冬成虫的耐寒性研究[J]. 山东农业科学, 2007(3):67-68.
[87] Denlinger D L. Regulation of diapause[J]. Annual Review of Entomology, 2001, 47(1):93-122.
[88] Willis J H. The Evolution and metamorphosis of arthropod proteomics and genomics[J]. Annual Review of Entomology, 2018, 63(1):1-13.
[89] Hahn D A, Denlinger D L. Energetics of insect diapause[J]. Annual Review of Entomology, 2011, 56(1):103-121.
[90] 蒋莎, 黄凤霞, 齐晓阳, 等. 滞育七星瓢虫抑制性消减杂交文库的构建及分析[J]. 中国生物防治学报, 2016, 32(1):33-39.
[91] Qi X Y, Zhang L S, Han Y H, et al. De novo transcriptome sequencing and analysis of Coccinella septempunctata L. in non-diapause, diapause and diapause-terminated states to identify diapause-associated genes[J]. BMC Genomics, 2015, 16(1):1086.
[92] 韩艳华, 刘梦姚, 高飞, 等. 滞育七星瓢虫酰基辅酶AΔ11去饱和酶基因的克隆及表达分析[J]. 中国生物防治学报, 2018, 34(5):31-40.
[93] Zhang H Z, Li Y Y, An T, et al. Comparative transcriptome and iTRAQ proteome analyses reveal the mechanisms of diapause in Aphidius gifuensis Ashmead (Hymenoptera:Aphidiidae)[J]. Frontiers in Physiology, 2018, 9:1697.
[94] Colinet H, Renault D, Charoy-Guevel B, et al. Metabolic and proteomic profiling of diapause in the aphid parasitoid Praon volucre[J]. PLoS ONE, 2012, 7(2):e32606.
[95] Li Y Y, Zhang L S, Chen H Y, et al. Shifts in metabolomic profiles of the parasitoid Nasonia vitripennis associated with elevated cold tolerance induced by the parasitoid's diapause, host diapause and host diet augmented with proline[J]. Insect Biochemistry and Molecular Biology, 2015, 63:34-46.
[96] Li Z Q, Zhang S, Ma Y, et al. First transcriptome and digital gene expression analysis in Neuroptera with an emphasis on chemoreception genes in Chrysopa pallens (Rambur)[J]. PLoS ONE, 2013, 8(6):e67151.
[97] Li Z Q, Zhang S, Cai X M, et al. Three odorant binding proteins may regulate the behavioural response of Chrysopa pallens to plant volatiles and the aphid alarm pheromone (E)-β-farnesene[J]. Insect Molecular Biology, 2017, 26(3):255-265.
[98] Zhang T T, Zhang G C, Zeng F F, et al. Insulin-like peptides regulate vitellogenesis and oviposition in the green lacewing, Chrysopa septempunctata[J]. Bulletin of Entomological Research, 2017, 107(2):148-154.
[99] Liu C Y, Mao J J, Zeng F R. Chrysopa septempunctata (Neuroptera:Chrysopidae) Vitellogenin functions through effects on egg production and hatching[J]. Journal of Economic Entomology, 2015, 108(6):2779.
[100] Han B F, Zhang S, Zeng F R, et al. Nutritional and reproductive signaling revealed by comparative gene expression analysis in Chrysopa pallens (Rambur) at different nutritional statuses[J]. PLoS ONE, 2017, 12(7):e0180373.
[101] 刘力源. 光周期调控下中华通草蛉滞育发生的基因表达谱分析及其对子代发育的影响[D]. 济南:山东农业大学, 2017.
[102] 王小平, 薛芳森. 昆虫滞育后的生物学特性[J]. 应用昆虫学报, 2006, 43(1):10-15.
[103] Danks H V. Insect adaptations to cold and changing environments[J]. Canadian Entomology, 2006, 138(1):1-23.
[104] 程丽媛, 张艳, 陈珍珍, 等. 光周期和温度对大草蛉滞育解除及滞育后发育和繁殖的影响[J]. 昆虫学报, 2017, 60(3):318-327.
[105] Beck S D. Insect Photoperiododism (2nd ed)[M]. New York:Academic Press, 1980, 288.
[106] Tauber M J, Tauber C A, Gardescu S. Prolonged storage of Chrysoperla carnea (Neuroptera:Chrysopidae)[J]. Environmental Entomology, 1993, 22(4):843-848.
[107] Kurbanov G G. Some data on mass laboratory rearing of the lacewings Chrysopa carnea Steph. and Ch. septempunctata Wesm[J]. Izvestiya Akademii Nauk AzerbaidzhanskoiSSR, Biologicheskikh Nauk, 1984(3):31-39.
[108] Chang Y F, Tauber M J, Tauber C A. Reproduction and quality of F1, offspring in Chrysoperla carnea:differential influence of quiescence, artificially-induced diapause, and natural diapause[J]. Journal of Insect Physiology, 1996, 42(6):521-528.
[109] Muntyan E M, Sekirov I A, Karelin V D, et al. Improving mass rearing of Chrysopa carnea[J]. Zashchita i Karantin Rastenii, 2012(6):21-22.
[110] Canard M. The possibilites of long-term preservation of the cocoons of a predator of aphids Chrysopa perla (L) (Neuroptera:Chrysopidae)[J]. Annales de Zoologie, Ecologie Animale, 1971, 3(3):373-377.