中华稻蝗不同龄期的形态多样性研究
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摘要
【目的】中华稻蝗Oxya chinensis的个体发育中经历体型渐变发育过程,随着龄期增加,一些功能器官逐渐发育完善,并执行相应的功能。研究中华稻蝗的个体发育规律,对于充分认识蝗虫体型的形成规律,机体结构与功能的关系,具有重要意义。【方法】本研究运用几何形态测量学方法分析不同虫龄的中华稻蝗身体结构的发育过程,探讨各器官结构的形成规律以及其功能的实现。【结果】研究结果表明:中华稻蝗不同龄的后翅、前胸背板、腹板、颜面、后足股节大小存在显著差异,随着虫龄的增加,各结构呈线性增大。PCA法和薄板样条法结果表明后翅从4龄到6龄,翅型结构呈渐变规律,成虫时肘脉域变小,轭脉域及臀脉域扩大,后翅的形状随其功能的实现显著不同于翅芽时期。前胸背板、颜面和后足股节随虫龄的增加在PC1轴呈渐变规律,随着取食和运动能力的增强,前胸背板变得更加宽阔、颜面的颊侧更加宽大,后足股节更加细长。腹板的形状在低龄阶段差异不大,成虫时在后足股节的着生区大而且向外突出。【结论】中华稻蝗不同虫龄间在后翅、前胸背板、腹板、颜面、后足股节等功能器官上存在显著差异。随着龄期的增加,各功能器官的形态呈渐变规律,然而这些功能器官的形成并非等比例扩大,而是形成特定的形态来实现其功能。
[Objectives] Oxya chinensis undergoes gradual morphological change during development and investigating these changes are important to understanding the form and function of adult grasshopper morphology. [Methods] The body structure of O. chinensis of different ages was analyzed using geometric morphometric methods and the regularity of the structure of each organ, and its functionality, were inferred. [Results] There were significant differences in the size of the hindwings, pronotum, sternum, frons and femur of O. chinensis of different ages. The size of these organs increased almost linearly with age. PCA and thin-plate spline analyses show that hindwing shape changed gradually from the fourth instar to the sixth; its cubital region becoming smaller, and the jugal and anal region expanding. The shape of the pronotum, frons and femur also changed gradually with age on the PC1 axis. Thin-plate spline results show that the pronotum became wider, the cheek of the frons broader and the femur slenderer. The adult and larval sternum do not differ significantly in shape, however the adult sternum is larger and has a more prominent femoral attachment area. [Conclusion] Various functional organs of O. chinensis, such as the hindwings, pronotum, sternum, frons and femur, differ significantly in shape in different developmental stages. The shape of these functional organs changed gradually with age, but, reflecting the functional requirements of each organ, different parts of each organs did not increase equally in size.
[Objectives] Oxya chinensis undergoes gradual morphological change during development and investigating these changes are important to understanding the form and function of adult grasshopper morphology. [Methods] The body structure of O. chinensis of different ages was analyzed using geometric morphometric methods and the regularity of the structure of each organ, and its functionality, were inferred. [Results] There were significant differences in the size of the hindwings, pronotum, sternum, frons and femur of O. chinensis of different ages. The size of these organs increased almost linearly with age. PCA and thin-plate spline analyses show that hindwing shape changed gradually from the fourth instar to the sixth; its cubital region becoming smaller, and the jugal and anal region expanding. The shape of the pronotum, frons and femur also changed gradually with age on the PC1 axis. Thin-plate spline results show that the pronotum became wider, the cheek of the frons broader and the femur slenderer. The adult and larval sternum do not differ significantly in shape, however the adult sternum is larger and has a more prominent femoral attachment area. [Conclusion] Various functional organs of O. chinensis, such as the hindwings, pronotum, sternum, frons and femur, differ significantly in shape in different developmental stages. The shape of these functional organs changed gradually with age, but, reflecting the functional requirements of each organ, different parts of each organs did not increase equally in size.
引文
Bai Y,Dong JJ,Guan DL,Xie JY,Xu SQ,2016.Geographic variation in wing size and shape of the grasshopper Trilophidia annulata(Orthoptera:Oedipodidae):morphological trait variations follow an ecogeographical rule.Scientific Reports,6:32680.DOI:10.1038/srep32680.
Bai M,Yang XK,Li J,Wang WC,2014.Geometric morphometrics,a super scientific computing tool in morphology comparison.Chinese Science Bulletin,59(10):887-894.[白明,杨星科,李静,王文成,2014.几何形态学:关于形态定量比较的科学计算工具.科学通报,59(10):887-894.]
Bookstein FL,1997.Landmark methods for forms without landmarks:morphometrics of group differences in outline shape.Medical Image Analysis,1(3):225-243.
Bowler K,Terblanche JS,2008.Insect thermal tolerance:what is the role of ontogeny,ageing and senescence?Biological Reviews,83(3):339-355.
Buckley LB,Arakaki AJ,Cannistra AF,Kharouba HM,Kingsolver JG,2017.Insect development,thermal plasticity and fitness implications in changing,seasonal environments.Integrative and Comparative Biology,57(5):988-998.
Cui SS,Zhu DH,2011.Embryonic development and the stages of diapause incidence in the Chinese rice grasshopper,Oxya chinensis.Chinese Journal of Applied Entomology,48(4):845-853.[崔双双,朱道弘,2011.中华稻蝗的胚胎发育及卵滞育发生的胚胎发育阶段.应用昆虫学报,48(4):845-853.]
Ding GW,Yu RR,Yang ML,Ma EB,Yang J,2014.Molecular characterization and functional analysis of chitin deacetylase 1gene in Oxya chinensis(Orthoptera:Acrididae).Acta Entomologica Sinica,57(11):1265-1271.[丁国伟,于荣荣,杨美玲,马恩波,杨静,2014.中华稻蝗几丁质脱乙酰基酶1基因的分子特性及功能.昆虫学报,57(11):1265-1271.]
Foth C,Ezcurra MD,Sookias RB,Brusatte SL,Butler RJ,2016.Unappreciated diversification of stem archosaurs during the middle triassic predated the dominance of dinosaurs.BMCEvolutionary Biology,16(1):188.https://doi.org/10.1186/s12862-016-0761-6.
Fusco G,Minelli A,2010.Phenotypic plasticity in development and evolution:facts and concepts.Philosophical Transactions of the Royal Society B,365(1540):547-556.
Gunz P,Mitteroecker P,2013.Semi-landmarks:a method for quantifying curves and surfaces.Hystrix,24(1):103-109.
Jockusch EL,Nulsen C,Newfeld SJ,Nagy LM,2000.Leg development in flies versus grasshoppers:differences in dpp expression do not lead to differences in the expression of downstream components of the leg patterning pathway.Development,127(8):1617-1626.
Klingenberg CP,Badyaev AV,Sowry SM,Beckwith NJ,2001.Inferring developmental modularity from morphological integration:analysis of individual variation and asymmetry in bumblebee wings.American Naturalist,157(1):11-23.
Klingenberg CP,2009.Morphometric integration and modularity in configurations of landmarks:tools for evaluating a priori hypotheses.Evolution&Development,11(4):405-421.
Klingenberg CP,Debat V,Roff DA,2010.Quantitative genetics of shape in cricket wings:developmental integration in a functional structure.Evolution,64(10):2935-2951.
Klingenberg CP,Gidaszewski NA,2010.Testing and quantifying phylogenetic signals and homoplasy in morphometric data.Systematic Biology,59(3):245-261.
Liu W,Clarke R,Xing S,2010.Geometric morphometric analysis of the early Pleistocene hominin teeth from Jianshi,Hubei Province,China.Science China Earth Sciences,40(6):724-736.[刘武,Clarke R,邢松,2010.湖北建始更新世早期人类牙齿几何形态测量分析.中国科学:地球科学,40(6):724-736.]
Louiseroth V,Mercer JM,2000.Morphometrics in development and evolution.American Zoologist,40(5):801-810.
Mayer C,Metscher BD,Müller GB,Mitteroecker P,2014.Studying developmental variation with geometric morphometric image analysis(GMIA).PLo S ONE,9(12):e115076.DOI:10.1371/journal.pone.0115076.
Monaghan P,2008.Early growth conditions,phenotypic development and environmental change.Philosophical Transactions of the Royal Society B,363(1497):1635-1645.
Ortego J,Aguirre MP,Cordero PJ,2011.Fine-scale spatial genetic structure and within population male-biased gene-flow in the grasshopper Mioscirtus wagneri.Evolutionary Ecology,25(5):1127-1144.
Régnière J,Powell J,Bentz B,Nealis V,2012.Effects of temperature on development,survival and reproduction of insects:experimental design,data analysis and modeling.Journal of Insect Physiology,58(5):634-647.
Ren LX,Yu ZT,Liu XJ,Zhang JZ,Ma EB,Guo YP,2014.Molecular characterization and expression analysis of metal responsive transcription factor1 gene Oc MTF1 from Oxya chinensis(Orthoptera:Acridoidea).Acta Entomologica Sinica,57(9):1001-1007.[任丽娴,余志涛,刘晓健,张建珍,马恩波,郭亚平,2014.中华稻蝗金属应答转录因子基因Oc MTF1的分子特性及表达分析.昆虫学报,57(9):1001-1007.]
Rohlf FJ,2006.Tps Dig2,version 2.10.Department of ecology and evolution,State University of New York,Stony Brook,NY.Available at http://life.bio.sunysb.edu/morph.
Rohlf FJ,Slice DE,1990.Extensions of the procrustes method for the optimal superimposition of landmarks.Systematic Zoology,39(1):40-59.
Scholtz G,Edgecombe GD,2006.The evolution of arthropod heads:reconciling morphological,developmental and palaeontological evidence.Development Genes and Evolution,216(7):395-415.
Sheets HD,2003.IMP-integrated morphometrics package.Department of Physics,Canisius College,Buffalo,New York.http://www.canisius.edu/~sheets/morphsoft.html.
Wang HD,Xu ZH,Feng ZQ,Xu FS,Wu YX,2007.The occurrence,damage and action of Chinese rice grasshopper,Oxya chinensis,in rice field.Acta Phytophylacic.Sinica,34(3):235-240.[王华弟,徐志宏,冯志全,徐福寿,吴玉香,2007.稻田中华稻蝗发生动态、危害损失及防治指标.植物保护学报,34(3):235-240.]
Wang SY,Wang GM,Yang ML,1993.The age recognition of Oxya.Chinese Journal of Applied Entomology,38(4):243-245.[王绍英,王光明,杨木林,1993.中华稻蝗蝗蝻龄期识别.应用昆虫学报,38(4):243-245.]
Wen GH,Bai Y,Zhou J,Zhang R,Zhang XX,Luo QY,Xu SQ,2015.Geometric morphometric analysis of wing shape variation in five Oxya spp.grasshoppers.Chinese Journal of Applied Entomology,52(2):356-362.[温光华,白义,周娟,张瑞,张秀秀,罗倩芸,许升全,2015.基于几何形态测量学的五种稻蝗前后翅的形态变化研究.应用昆虫学报,52(2):356-362.]
Wootton RJ,Evans KE,Herbert R,Smith CW,2000.The hind wing of the desert locust(Schistocerca gregaria Forsk?l).Journal of Experimental Biology,203(19):2921-2931.
Zhang HW,2013.Developmental Biology.3rd ed.Beijing:Higher Education Press.1-321.[张红卫,2013.发育生物学(第3版).北京:高等教育出版社.1-321.]
Zhao HQ,Shen ZR,Yu XW,2003.Use of math-morphological features in insect taxonomy.I.At the order level.Acta Entomologica Sinica,46(1):45-50.[赵汗青,沈佐锐,于新闻,2003.数学形态学在昆虫分类学上的应用研究.I.在目级阶元上的应用研究.昆虫学报,46(1):45-50.]
Zheng ZM,Xu WX,Lian ZM,1990.Grasshoppers in Shaanxi.Shaanxi:Shaanxi Normal University Press.18-21.[郑哲民,许文贤,廉振民,1990.陕西蝗虫.陕西:陕西师范大学出版社.18-21.]
Bai M,Yang XK,Li J,Wang WC,2014.Geometric morphometrics,a super scientific computing tool in morphology comparison.Chinese Science Bulletin,59(10):887-894.[白明,杨星科,李静,王文成,2014.几何形态学:关于形态定量比较的科学计算工具.科学通报,59(10):887-894.]
Bookstein FL,1997.Landmark methods for forms without landmarks:morphometrics of group differences in outline shape.Medical Image Analysis,1(3):225-243.
Bowler K,Terblanche JS,2008.Insect thermal tolerance:what is the role of ontogeny,ageing and senescence?Biological Reviews,83(3):339-355.
Buckley LB,Arakaki AJ,Cannistra AF,Kharouba HM,Kingsolver JG,2017.Insect development,thermal plasticity and fitness implications in changing,seasonal environments.Integrative and Comparative Biology,57(5):988-998.
Cui SS,Zhu DH,2011.Embryonic development and the stages of diapause incidence in the Chinese rice grasshopper,Oxya chinensis.Chinese Journal of Applied Entomology,48(4):845-853.[崔双双,朱道弘,2011.中华稻蝗的胚胎发育及卵滞育发生的胚胎发育阶段.应用昆虫学报,48(4):845-853.]
Ding GW,Yu RR,Yang ML,Ma EB,Yang J,2014.Molecular characterization and functional analysis of chitin deacetylase 1gene in Oxya chinensis(Orthoptera:Acrididae).Acta Entomologica Sinica,57(11):1265-1271.[丁国伟,于荣荣,杨美玲,马恩波,杨静,2014.中华稻蝗几丁质脱乙酰基酶1基因的分子特性及功能.昆虫学报,57(11):1265-1271.]
Foth C,Ezcurra MD,Sookias RB,Brusatte SL,Butler RJ,2016.Unappreciated diversification of stem archosaurs during the middle triassic predated the dominance of dinosaurs.BMCEvolutionary Biology,16(1):188.https://doi.org/10.1186/s12862-016-0761-6.
Fusco G,Minelli A,2010.Phenotypic plasticity in development and evolution:facts and concepts.Philosophical Transactions of the Royal Society B,365(1540):547-556.
Gunz P,Mitteroecker P,2013.Semi-landmarks:a method for quantifying curves and surfaces.Hystrix,24(1):103-109.
Jockusch EL,Nulsen C,Newfeld SJ,Nagy LM,2000.Leg development in flies versus grasshoppers:differences in dpp expression do not lead to differences in the expression of downstream components of the leg patterning pathway.Development,127(8):1617-1626.
Klingenberg CP,Badyaev AV,Sowry SM,Beckwith NJ,2001.Inferring developmental modularity from morphological integration:analysis of individual variation and asymmetry in bumblebee wings.American Naturalist,157(1):11-23.
Klingenberg CP,2009.Morphometric integration and modularity in configurations of landmarks:tools for evaluating a priori hypotheses.Evolution&Development,11(4):405-421.
Klingenberg CP,Debat V,Roff DA,2010.Quantitative genetics of shape in cricket wings:developmental integration in a functional structure.Evolution,64(10):2935-2951.
Klingenberg CP,Gidaszewski NA,2010.Testing and quantifying phylogenetic signals and homoplasy in morphometric data.Systematic Biology,59(3):245-261.
Liu W,Clarke R,Xing S,2010.Geometric morphometric analysis of the early Pleistocene hominin teeth from Jianshi,Hubei Province,China.Science China Earth Sciences,40(6):724-736.[刘武,Clarke R,邢松,2010.湖北建始更新世早期人类牙齿几何形态测量分析.中国科学:地球科学,40(6):724-736.]
Louiseroth V,Mercer JM,2000.Morphometrics in development and evolution.American Zoologist,40(5):801-810.
Mayer C,Metscher BD,Müller GB,Mitteroecker P,2014.Studying developmental variation with geometric morphometric image analysis(GMIA).PLo S ONE,9(12):e115076.DOI:10.1371/journal.pone.0115076.
Monaghan P,2008.Early growth conditions,phenotypic development and environmental change.Philosophical Transactions of the Royal Society B,363(1497):1635-1645.
Ortego J,Aguirre MP,Cordero PJ,2011.Fine-scale spatial genetic structure and within population male-biased gene-flow in the grasshopper Mioscirtus wagneri.Evolutionary Ecology,25(5):1127-1144.
Régnière J,Powell J,Bentz B,Nealis V,2012.Effects of temperature on development,survival and reproduction of insects:experimental design,data analysis and modeling.Journal of Insect Physiology,58(5):634-647.
Ren LX,Yu ZT,Liu XJ,Zhang JZ,Ma EB,Guo YP,2014.Molecular characterization and expression analysis of metal responsive transcription factor1 gene Oc MTF1 from Oxya chinensis(Orthoptera:Acridoidea).Acta Entomologica Sinica,57(9):1001-1007.[任丽娴,余志涛,刘晓健,张建珍,马恩波,郭亚平,2014.中华稻蝗金属应答转录因子基因Oc MTF1的分子特性及表达分析.昆虫学报,57(9):1001-1007.]
Rohlf FJ,2006.Tps Dig2,version 2.10.Department of ecology and evolution,State University of New York,Stony Brook,NY.Available at http://life.bio.sunysb.edu/morph.
Rohlf FJ,Slice DE,1990.Extensions of the procrustes method for the optimal superimposition of landmarks.Systematic Zoology,39(1):40-59.
Scholtz G,Edgecombe GD,2006.The evolution of arthropod heads:reconciling morphological,developmental and palaeontological evidence.Development Genes and Evolution,216(7):395-415.
Sheets HD,2003.IMP-integrated morphometrics package.Department of Physics,Canisius College,Buffalo,New York.http://www.canisius.edu/~sheets/morphsoft.html.
Wang HD,Xu ZH,Feng ZQ,Xu FS,Wu YX,2007.The occurrence,damage and action of Chinese rice grasshopper,Oxya chinensis,in rice field.Acta Phytophylacic.Sinica,34(3):235-240.[王华弟,徐志宏,冯志全,徐福寿,吴玉香,2007.稻田中华稻蝗发生动态、危害损失及防治指标.植物保护学报,34(3):235-240.]
Wang SY,Wang GM,Yang ML,1993.The age recognition of Oxya.Chinese Journal of Applied Entomology,38(4):243-245.[王绍英,王光明,杨木林,1993.中华稻蝗蝗蝻龄期识别.应用昆虫学报,38(4):243-245.]
Wen GH,Bai Y,Zhou J,Zhang R,Zhang XX,Luo QY,Xu SQ,2015.Geometric morphometric analysis of wing shape variation in five Oxya spp.grasshoppers.Chinese Journal of Applied Entomology,52(2):356-362.[温光华,白义,周娟,张瑞,张秀秀,罗倩芸,许升全,2015.基于几何形态测量学的五种稻蝗前后翅的形态变化研究.应用昆虫学报,52(2):356-362.]
Wootton RJ,Evans KE,Herbert R,Smith CW,2000.The hind wing of the desert locust(Schistocerca gregaria Forsk?l).Journal of Experimental Biology,203(19):2921-2931.
Zhang HW,2013.Developmental Biology.3rd ed.Beijing:Higher Education Press.1-321.[张红卫,2013.发育生物学(第3版).北京:高等教育出版社.1-321.]
Zhao HQ,Shen ZR,Yu XW,2003.Use of math-morphological features in insect taxonomy.I.At the order level.Acta Entomologica Sinica,46(1):45-50.[赵汗青,沈佐锐,于新闻,2003.数学形态学在昆虫分类学上的应用研究.I.在目级阶元上的应用研究.昆虫学报,46(1):45-50.]
Zheng ZM,Xu WX,Lian ZM,1990.Grasshoppers in Shaanxi.Shaanxi:Shaanxi Normal University Press.18-21.[郑哲民,许文贤,廉振民,1990.陕西蝗虫.陕西:陕西师范大学出版社.18-21.]
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