淡黄蚊蝎蛉染色体特征及其系统发育意义
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摘要
【目的】染色体特征在昆虫系统发育研究中起着重要作用。然而,长翅目(Mecoptera)蚊蝎蛉科(Bittacidae)昆虫的染色体研究却比较匮乏。【方法】通过室内饲养获得淡黄蚊蝎Bittacus flavidus Huang&Hua各龄幼虫、蛹和成虫。取淡黄蚊蝎雄性末龄(4龄)幼虫、蛹和新羽化成虫精巢细胞进行染色体制片,通过4′,6-二脒基-2-苯基吲哚(4′,6-diamidino-2-phenylindole, DAPI)荧光染色,研究染色体核型、减数分裂行为和性别决定机制。【结果】结果表明,淡黄蚊蝎蛉的染色体数目为2n=26+X0,染色体均具有中着丝粒,核型高度对称。二价体绝对长度为(3.20±0.07)～(1.53±0.19)μm,相对长度为(5.31±0.29)～(2.73±0.24),均呈梯度变化。淡黄蚊蝎蛉的减数分裂为交叉型,其中核的平均交叉频率为11.5,二价体的平均交叉频率为0.88。性别决定机制为XX/X0型。DAPI荧光带型显示,粗线期二价体一端呈现高亮的AT富集区块。【结论】染色体特征(包括染色体数目、染色体臂基本数和核型公式等)在蚊蝎蛉科中表现出显著变异,表明染色体重排(尤其是融合、断裂和倒位)在蚊蝎蛉科昆虫的谱系分化和染色体演化中起着重要作用。
【Aim】 Chromosomal characteristics play significant roles in phylogenetic analyses of insects. However, chromosomes in Bittacidae(Mecoptera) have rarely been studied to date. 【Methods】 Larvae, pupae and adults of the hangingfly Bittacus flavidus Huang & Hua were obtained by rearing in the laboratory. Chromosome preparations for the testes of the last(4 th) instar larvae, pupae and newly-emerged adults of the hangingfly were stained with 4′,6-diamidino-2-phenylindole(DAPI) to investigate the karyotype, meiotic behavior, and sex determination. 【Results】 The results showed that the chromosome number of B. flavidus is 2 n=26+X0, and all the chromosomes are metacentric, showing a symmetric karyotype. The absolute length of bivalents ranged from 3.20±0.07 to 1.53±0.19 μm, and the relative length from 5.31±0.29 to 2.73±0.24, with their size decreasing gradually from pair to pair. The meiosis of B. flavidus is chiasmate with a mean chiasma frequency of 11.5 per nucleus and a mean chiasma frequency of 0.88 per autosomal bivalent. The sex determination mechanism is of the XX/X0 type. Fluorescent bands with DAPI staining revealed a large AT-rich block covering one terminal region on all pachytene bivalents. 【Conclusion】 The chromosomes exhibit marked variations with respect to the diploid number, fundamental number and karyotype formula, suggesting that chromosome rearrangements(especially fusion, fission and inversion) play an important part in the lineage differentiation and chromosome evolution in Bittacidae.
【Aim】 Chromosomal characteristics play significant roles in phylogenetic analyses of insects. However, chromosomes in Bittacidae(Mecoptera) have rarely been studied to date. 【Methods】 Larvae, pupae and adults of the hangingfly Bittacus flavidus Huang & Hua were obtained by rearing in the laboratory. Chromosome preparations for the testes of the last(4 th) instar larvae, pupae and newly-emerged adults of the hangingfly were stained with 4′,6-diamidino-2-phenylindole(DAPI) to investigate the karyotype, meiotic behavior, and sex determination. 【Results】 The results showed that the chromosome number of B. flavidus is 2 n=26+X0, and all the chromosomes are metacentric, showing a symmetric karyotype. The absolute length of bivalents ranged from 3.20±0.07 to 1.53±0.19 μm, and the relative length from 5.31±0.29 to 2.73±0.24, with their size decreasing gradually from pair to pair. The meiosis of B. flavidus is chiasmate with a mean chiasma frequency of 11.5 per nucleus and a mean chiasma frequency of 0.88 per autosomal bivalent. The sex determination mechanism is of the XX/X0 type. Fluorescent bands with DAPI staining revealed a large AT-rich block covering one terminal region on all pachytene bivalents. 【Conclusion】 The chromosomes exhibit marked variations with respect to the diploid number, fundamental number and karyotype formula, suggesting that chromosome rearrangements(especially fusion, fission and inversion) play an important part in the lineage differentiation and chromosome evolution in Bittacidae.
引文
Atchley WR,Jackson RC,1970.Cytological observations on spermatogenesis in four species of Mecoptera.Can.J.Genet.Cytol.,12(2):264-272.
Beutel RG,Baum E,2008.A longstanding entomological problem finally solved?Head morphology of Nannochorista (Mecoptera,Insecta) and possible phylogenetic implications.J.Zool.Syst.Evol.Res.,46(4):346-367.
Beutel RG,Friedrich F,H?rnschemeyer T,Pohl H,Hünefeld F,Beckmann F,Meier R,Misof B,Whiting MF,Vilhelmsen L,2011.Morphological and molecular evidence converge upon a robust phylogeny of the megadiverse Holometabola.Cladistics,27(4):341-355.
Bickmore WA,2001.Karyotype analysis and chromosome banding.In:Cooper DN ed.Encyclopedia of Life Sciences.John Wiley & Sons,Chichester.1-7.
Bidau CJ,Mirol PM,1988.Orientation and segregation of Robertsonian trivalents in Dichroplus pratensis (Acrididae).Genome,30(6):947-955.
Bornemissza GF,1966.Observations on the hunting and mating behaviours of two species of scorpion flies (Bittacidae:Mecoptera).Aust.J.Zool.,14(3):371-382.
Bush GL,1967.The comparative cytology of the Choristidae and Nannochoristidae (Mecoptera).Am.Philos.Soc.Yearb.,1966:326-328.
Cabral-de-Mello DC,Oliveira SG,de Moura RC,Martins C,2011.Chromosomal organization of the 18S and 5S rRNAs and histone H3 genes in Scarabaeinae coleopterans:insights into the evolutionary dynamics of multigene families and heterochromatin.BMC Genetics,12:88.
Castillo ER,Marti DA,Bidau CJ,2010.Sex and neo-sex chromosomes in Orthoptera:a review.J.Orthopt.Res.,19(2):213-231.
Castillo ERD,Martí DA,Maronna MM,Scattolini MC,Cabral-de-Mello DC,Cigliano MM,2019.Chromosome evolution and phylogeny in Ronderosia (Orthoptera,Acrididae,Melanoplinae):clues of survivors to the challenge of sympatry?Syst.Entomol.,44(1):61-74.
Cooper KW,1951.Compound sex chromosomes with anaphasic precocity in the male mecopteran,Boreus brumalis Fitch.J.Morphol.,89(1):37-57.
Cooper KW,1974.Sexual biology,chromosomes,development,life histories and parasites of Boreus,especially of B.notoperates.A southern California Boreus.Ⅱ.(Mecoptera:Boreidae).Psyche,81:84-120.
Dallai R,Lupetti P,Afzelius BA,Frati F,2003.Sperm structure of Mecoptera and Siphonaptera (Insecta) and the phylogenetic position of Boreus hyemalis.Zoomorphology,122(4):211-220.
Dobigny G,Ducroz JF,Robinson TJ,Volobouev V,2004.Cytogenetics and cladistics.Syst.Biol.,53(3):470-484.
Dyer AF,1979.Investigating Chromosomes.Edward Arnold,London.
Faria R,Navarro A,2010.Chromosomal speciation revisited:rearranging theory with pieces of evidence.Trends Ecol.Evol.,25(11):660-669.
Friedrich F,Pohl H,Beckmann F,Beutel RG,2013.The head of Merope tuber (Meropeidae) and the phylogeny of Mecoptera (Hexapoda).Arthropod Struct.Dev.,42(1):69-88.
Gao QH,Hua BZ,2013.Co-evolution of the mating position and male genitalia in insects:a case study of a hangingfly.PLoS ONE,8(12):e80651.
Gokhman VE,2009.Karyotypes of Parasitic Hymenoptera.Springer Science & Business Media B.V.,Dordrecht.
Gokhman VE,Kuznetsova VG,2006.Comparative insect karyology:current state and applications.Entomol.Rev.,86(3):352-368.
Hughes-Schrader S,1950.The chromosomes of mantids (Orthoptera:Manteidae) in relation to taxonomy.Chromosoma,4(1):1-55.
Imai HT,Taylor RW,Crosland MW,Crozier RH,1988.Modes of spontaneous chromosomal mutation and karyotype evolution in ants with reference to the minimum interaction hypothesis.Jpn.J.Genet.,63(2):159-185.
Jiang L,Gao QH,Hua BZ,2015.Larval morphology of the hanging-fly Bittacus trapezoideus Huang & Hua (Insecta:Mecoptera:Bittacidae).Zootaxa,3957(3):324-333.
Lambkin KJ,1988.An Australian species of the genus Bittacus Latreille (Mecoptera:Bittacidae).Mem.Queensland Mus.,25(2):439-444.
Levan A,Fredga K,Sandberg AA,1964.Nomenclature for centromeric position on chromosomes.Hereditas,52(2):201-220.
Lukhtanov VA,Dinc V,Talavera G,Vila R,2011.Unprecedented within-species chromosome number cline in the wood white butterfly Leptidea sinapis and its significance for karyotype evolution and speciation.BMC Evol.Biol.,11:109.
Lukhtanov VA,Kandul NP,Plotkin JB,Dantchenko AV,Haig D,Pierce NE,2005.Reinforcement of pre-zygotic isolation and karyotype evolution in Agrodiaetus butterflies.Nature,436(7049):385-389.
Ma N,Huang J,Hua BZ,2014.Fine structure and functional morphology of the mouthparts of Bittacus planus and Terrobittacus implicatus (Insecta:Mecoptera:Bittacidae).Zool.Anz.,253(6):441-448.
Matthey R,1950.La formule chromosomique et le type de digamétie chez Bittacus italicus Müll.(Mecoptera).Arch.Klaus-Stift.Vererb.Forsch.,25:605-611.
McAllister BF,Sheeley SL,Mena PA,Evans AL,Schl?tterer C,2008.Clinal distribution of a chromosomal rearrangement:a precursor to chromosomal speciation?Evolution,62(8):1852-1865.
Miao Y,Hua BZ,2017.Cytogenetic comparison between Terrobittacus implicatus and Bittacus planus (Mecoptera:Bittacidae) with some phylogenetic implications.Arthropod Syst.Phyl.,75(2):175-183.
Miao Y,Ma N,Hua BZ,2017.Cytotaxonomy and molecular phylogeny of the genus Cerapanorpa Gao,Ma & Hua,2016 (Mecoptera:Panorpidae).Sci.Rep.,7:4493.
Miao Y,Wang JS,Hua BZ,2019.Molecular phylogeny of the scorpionflies Panorpidae (Insecta:Mecoptera) and chromosomal evolution.Cladistics,https://doi.org/10.1111/cla.12357.
Mickoleit G,Mickoleit E,1978.Zum Kopulationsverhalten des Mückenhaftes Bittacus italicus (Mecoptera:Bittacidae).Entomol.Gen.,5:1-15.
Mills PJ,Cook LG,2014.Rapid chromosomal evolution in a morphologically cryptic radiation.Mol.Phylogenet.Evol.,77:126-135.
Misof B,Liu SL,Meusemann K,et al.,2014.Phylogenomics resolves the timing and pattern of insect evolution.Science,346(6210):763-767.
Naville A,Beaumont J,1934.Les chromosomes des Panorpes.Bull.Biol.France Belg.,68:98-107.
Paszko B,2006.A critical review and a new proposal of karyotype asymmetry indices.Plant Syst.Evol.,258(1):39-48.
Penny ND,1975.Evolution of the extant Mecoptera.J.Kans.Entomol.Soc.,48(3):331-350.
Penny ND,Byers GW,1979.Check-list of the Mecoptera of the world.Acta Amazon.,9:365-388.
Rebagliati PJ,Papeschi AG,Mola LM,2003.Meiosis and fluorescent banding in Edessa meditabunda and E.rufomarginata (Heteroptera:Pentatomidae:Edessinae).Eur.J.Entomol.,100(1):11-18.
Stebbins GL,1971.Chromosomal Evolution in Higher Plants.Edward Arnold,London.
Tan JL,Hua BZ,2008.Structure of raptorial legs in Bittacus (Mecoptera:Bittacidae).Acta Entomol.Sin.,51(7):745-752.[谭江丽,花保祯,2008.蚊蝎蛉捕捉足构造.昆虫学报,51(7):745-752]
Ullerich FH,1961.Achiasmatische Spermatogenese bei der Skorpionsfliege Panorpa (Mecoptera).Chromosoma,12(1):215-232.
White MJD,1965.Sex chromosomes and meiotic mechanisms in some African and Australian mantids.Chromosoma,16(5):521-547.
White MJD,1974.Genetic Mechanisms of Speciation in Insects.Springer,Dordrecht.
Whiting MF,2002.Mecoptera is paraphyletic:multiple genes and phylogeny of Mecoptera and Siphonaptera.Zool.Scr.,31(1):93-104.
Wiegmann BM,Trautwein MD,Kim JW,Cassel BK,Bertone MA,Winterton SL,Yeates DK,2009.Single-copy nuclear genes resolve the phylogeny of the holometabolous insects.BMC Biol.,7:34.
Willmann R,1987.The phylogenetic system of the Mecoptera.Syst.Entomol.,12(4):519-524.
Willmann R,1989.Evolution und phylogenetisches System der Mecoptera (Insecta:Holometabola).Abh.Senckenb.Natforsch.Ges.,544:1-153.
Xu B,Li YK,Hua BZ,2013.A chromosomal investigation of four species of Chinese Panorpidae (Insecta,Mecoptera).Comp.Cytogenet.,7(3):229-239.
Beutel RG,Baum E,2008.A longstanding entomological problem finally solved?Head morphology of Nannochorista (Mecoptera,Insecta) and possible phylogenetic implications.J.Zool.Syst.Evol.Res.,46(4):346-367.
Beutel RG,Friedrich F,H?rnschemeyer T,Pohl H,Hünefeld F,Beckmann F,Meier R,Misof B,Whiting MF,Vilhelmsen L,2011.Morphological and molecular evidence converge upon a robust phylogeny of the megadiverse Holometabola.Cladistics,27(4):341-355.
Bickmore WA,2001.Karyotype analysis and chromosome banding.In:Cooper DN ed.Encyclopedia of Life Sciences.John Wiley & Sons,Chichester.1-7.
Bidau CJ,Mirol PM,1988.Orientation and segregation of Robertsonian trivalents in Dichroplus pratensis (Acrididae).Genome,30(6):947-955.
Bornemissza GF,1966.Observations on the hunting and mating behaviours of two species of scorpion flies (Bittacidae:Mecoptera).Aust.J.Zool.,14(3):371-382.
Bush GL,1967.The comparative cytology of the Choristidae and Nannochoristidae (Mecoptera).Am.Philos.Soc.Yearb.,1966:326-328.
Cabral-de-Mello DC,Oliveira SG,de Moura RC,Martins C,2011.Chromosomal organization of the 18S and 5S rRNAs and histone H3 genes in Scarabaeinae coleopterans:insights into the evolutionary dynamics of multigene families and heterochromatin.BMC Genetics,12:88.
Castillo ER,Marti DA,Bidau CJ,2010.Sex and neo-sex chromosomes in Orthoptera:a review.J.Orthopt.Res.,19(2):213-231.
Castillo ERD,Martí DA,Maronna MM,Scattolini MC,Cabral-de-Mello DC,Cigliano MM,2019.Chromosome evolution and phylogeny in Ronderosia (Orthoptera,Acrididae,Melanoplinae):clues of survivors to the challenge of sympatry?Syst.Entomol.,44(1):61-74.
Cooper KW,1951.Compound sex chromosomes with anaphasic precocity in the male mecopteran,Boreus brumalis Fitch.J.Morphol.,89(1):37-57.
Cooper KW,1974.Sexual biology,chromosomes,development,life histories and parasites of Boreus,especially of B.notoperates.A southern California Boreus.Ⅱ.(Mecoptera:Boreidae).Psyche,81:84-120.
Dallai R,Lupetti P,Afzelius BA,Frati F,2003.Sperm structure of Mecoptera and Siphonaptera (Insecta) and the phylogenetic position of Boreus hyemalis.Zoomorphology,122(4):211-220.
Dobigny G,Ducroz JF,Robinson TJ,Volobouev V,2004.Cytogenetics and cladistics.Syst.Biol.,53(3):470-484.
Dyer AF,1979.Investigating Chromosomes.Edward Arnold,London.
Faria R,Navarro A,2010.Chromosomal speciation revisited:rearranging theory with pieces of evidence.Trends Ecol.Evol.,25(11):660-669.
Friedrich F,Pohl H,Beckmann F,Beutel RG,2013.The head of Merope tuber (Meropeidae) and the phylogeny of Mecoptera (Hexapoda).Arthropod Struct.Dev.,42(1):69-88.
Gao QH,Hua BZ,2013.Co-evolution of the mating position and male genitalia in insects:a case study of a hangingfly.PLoS ONE,8(12):e80651.
Gokhman VE,2009.Karyotypes of Parasitic Hymenoptera.Springer Science & Business Media B.V.,Dordrecht.
Gokhman VE,Kuznetsova VG,2006.Comparative insect karyology:current state and applications.Entomol.Rev.,86(3):352-368.
Hughes-Schrader S,1950.The chromosomes of mantids (Orthoptera:Manteidae) in relation to taxonomy.Chromosoma,4(1):1-55.
Imai HT,Taylor RW,Crosland MW,Crozier RH,1988.Modes of spontaneous chromosomal mutation and karyotype evolution in ants with reference to the minimum interaction hypothesis.Jpn.J.Genet.,63(2):159-185.
Jiang L,Gao QH,Hua BZ,2015.Larval morphology of the hanging-fly Bittacus trapezoideus Huang & Hua (Insecta:Mecoptera:Bittacidae).Zootaxa,3957(3):324-333.
Lambkin KJ,1988.An Australian species of the genus Bittacus Latreille (Mecoptera:Bittacidae).Mem.Queensland Mus.,25(2):439-444.
Levan A,Fredga K,Sandberg AA,1964.Nomenclature for centromeric position on chromosomes.Hereditas,52(2):201-220.
Lukhtanov VA,Dinc V,Talavera G,Vila R,2011.Unprecedented within-species chromosome number cline in the wood white butterfly Leptidea sinapis and its significance for karyotype evolution and speciation.BMC Evol.Biol.,11:109.
Lukhtanov VA,Kandul NP,Plotkin JB,Dantchenko AV,Haig D,Pierce NE,2005.Reinforcement of pre-zygotic isolation and karyotype evolution in Agrodiaetus butterflies.Nature,436(7049):385-389.
Ma N,Huang J,Hua BZ,2014.Fine structure and functional morphology of the mouthparts of Bittacus planus and Terrobittacus implicatus (Insecta:Mecoptera:Bittacidae).Zool.Anz.,253(6):441-448.
Matthey R,1950.La formule chromosomique et le type de digamétie chez Bittacus italicus Müll.(Mecoptera).Arch.Klaus-Stift.Vererb.Forsch.,25:605-611.
McAllister BF,Sheeley SL,Mena PA,Evans AL,Schl?tterer C,2008.Clinal distribution of a chromosomal rearrangement:a precursor to chromosomal speciation?Evolution,62(8):1852-1865.
Miao Y,Hua BZ,2017.Cytogenetic comparison between Terrobittacus implicatus and Bittacus planus (Mecoptera:Bittacidae) with some phylogenetic implications.Arthropod Syst.Phyl.,75(2):175-183.
Miao Y,Ma N,Hua BZ,2017.Cytotaxonomy and molecular phylogeny of the genus Cerapanorpa Gao,Ma & Hua,2016 (Mecoptera:Panorpidae).Sci.Rep.,7:4493.
Miao Y,Wang JS,Hua BZ,2019.Molecular phylogeny of the scorpionflies Panorpidae (Insecta:Mecoptera) and chromosomal evolution.Cladistics,https://doi.org/10.1111/cla.12357.
Mickoleit G,Mickoleit E,1978.Zum Kopulationsverhalten des Mückenhaftes Bittacus italicus (Mecoptera:Bittacidae).Entomol.Gen.,5:1-15.
Mills PJ,Cook LG,2014.Rapid chromosomal evolution in a morphologically cryptic radiation.Mol.Phylogenet.Evol.,77:126-135.
Misof B,Liu SL,Meusemann K,et al.,2014.Phylogenomics resolves the timing and pattern of insect evolution.Science,346(6210):763-767.
Naville A,Beaumont J,1934.Les chromosomes des Panorpes.Bull.Biol.France Belg.,68:98-107.
Paszko B,2006.A critical review and a new proposal of karyotype asymmetry indices.Plant Syst.Evol.,258(1):39-48.
Penny ND,1975.Evolution of the extant Mecoptera.J.Kans.Entomol.Soc.,48(3):331-350.
Penny ND,Byers GW,1979.Check-list of the Mecoptera of the world.Acta Amazon.,9:365-388.
Rebagliati PJ,Papeschi AG,Mola LM,2003.Meiosis and fluorescent banding in Edessa meditabunda and E.rufomarginata (Heteroptera:Pentatomidae:Edessinae).Eur.J.Entomol.,100(1):11-18.
Stebbins GL,1971.Chromosomal Evolution in Higher Plants.Edward Arnold,London.
Tan JL,Hua BZ,2008.Structure of raptorial legs in Bittacus (Mecoptera:Bittacidae).Acta Entomol.Sin.,51(7):745-752.[谭江丽,花保祯,2008.蚊蝎蛉捕捉足构造.昆虫学报,51(7):745-752]
Ullerich FH,1961.Achiasmatische Spermatogenese bei der Skorpionsfliege Panorpa (Mecoptera).Chromosoma,12(1):215-232.
White MJD,1965.Sex chromosomes and meiotic mechanisms in some African and Australian mantids.Chromosoma,16(5):521-547.
White MJD,1974.Genetic Mechanisms of Speciation in Insects.Springer,Dordrecht.
Whiting MF,2002.Mecoptera is paraphyletic:multiple genes and phylogeny of Mecoptera and Siphonaptera.Zool.Scr.,31(1):93-104.
Wiegmann BM,Trautwein MD,Kim JW,Cassel BK,Bertone MA,Winterton SL,Yeates DK,2009.Single-copy nuclear genes resolve the phylogeny of the holometabolous insects.BMC Biol.,7:34.
Willmann R,1987.The phylogenetic system of the Mecoptera.Syst.Entomol.,12(4):519-524.
Willmann R,1989.Evolution und phylogenetisches System der Mecoptera (Insecta:Holometabola).Abh.Senckenb.Natforsch.Ges.,544:1-153.
Xu B,Li YK,Hua BZ,2013.A chromosomal investigation of four species of Chinese Panorpidae (Insecta,Mecoptera).Comp.Cytogenet.,7(3):229-239.