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Grozeva Et Al.Indd © Comparative Cytogenetics, 2008 . Vol. 2, No. 1, P. 73-78. ISSN 1993-0771 (Print), ISSN 1993-078X (Online) Achiasmatic male meiosis in three Micronecta species (Heteroptera: Nepomorpha: Micronectidae) S. Grozeva1, N. Simov2, S. Nokkala3 1Institute of Zoology, Bulgarian Academy of Sciences, 1 Tsar Osvoboditel, Sofia 1000, Bulgaria. 2National Museum of Natural History, Bulgarian Academy of Sci- ences, 1 Tsar Osvoboditel, Sofia 1000, Bulgaria. 3Laboratory of Genetics, Depart- ment of Biology, University of Turku, Turku FIN-20014, Finland. E-mails: [email protected], [email protected], [email protected], [email protected] Abstract. Male meiosis of three water boatmen species Micronecta (Dichaetonec- ta) scholtzi (Fieber, 1860), M. (Micronecta) poweri (Douglas et Scott, 1869) and M. (Micronecta) griseola Horvath, 1899 (Corixoidea: Micronectidae) was studied pay- ing special attention to the presence or absence of chiasmata. Condensation stage revealed 11 autosomal bivalents consisting of side by side associated homologous chromosomes. At this stage positively heterochromatic X and Y chromosomes were attached to the nucleolus. There were no m-chromosomes. The chromosome number of all species is 2n=22+XY. Side by side association of homologous chromosomes was still seen at late condensation stage. At prometaphase and metaphase I, the bi- valents oriented parallel to the equatorial plane and homologous chromosomes are facing opposite poles. Hence, male meiosis in these species is achiasmatic of align- ment type. At metaphase II chromosomes showed variable positions in the plate. In M. scholtzi, the Y chromosome is smaller than the X chromosome, while in M. pow- eri and M. griseola X and Y chromosomes were similar in size. It is suggested that achiasmatic male meiosis and lack of m-chromosomes are cytogenetic features sepa- rating the family Micronectidae from the family Corixidae among the superfamily Corixoidea within the infraorder Nepomorpha. It seems plausible that achiasmatic male meiosis has emerged in Heteroptera more than once during their evolution. Key words: Heteroptera, Micronectidae, achiasmatic male meiosis. INTRODUCTION In Hemiptera, achiasmatic meiosis has been found both in Homoptera (Schrader, 1931; Most often during meiosis, chiasma for- Nur, 1965; Blackman, 1976; Hales, 1989) and mation ties homologous chromosomes to- in six families of Heteroptera, belonging to gether until their separation in the reductional two infraorders: Leptopodomorpha and Cimi- division. However, in some groups of animals comorpha (Nokkala, Nokkala, 1983; Nokkala, chiasma formation during meiosis is replaced Nokkala, 1984; Nokkala, Nokkala, 1986a, b; by other, achiasmatic means. Achiasmatic Nokkala, Grozeva, 2000; Grozeva, Nokkala, meiosis, as a rule, is restricted to the hetero- 2002). Recently, achiasmatic meiosis had been gametic sex only, and has been described in reported for the first time in the infraorder different groups of animals (White, 1973). Nepomorpha (Ituarte, Papeschi, 2004). Ac- Free access: www.zin.ru/journals/compcyt 74 S. Grozeva , N. Simov, S. Nokkala cording to previous cytogenetic reports on ta) griseola Horvath, 1899 was studied paying 28 species of family Corixidae (Corixoidea) special attention to the presence or absence of 27 species display pre-reductional chiasmatic chiasmata. meiosis and a stable chromosome formula 2n=24(20+2m+XY) (except in Cymatia bons- MATERIAL AND METHODS dorffi (Sahlberg, 1819), 2n=26) (Ituarte, Pape- A number of males of three Micronecta schi, 2004; Bressa, Papeschi, 2007). species – 49 males of Micronecta (Dichaeto- Fieber (1851) was the first who considered necta) scholtzi, 6 males of M. (Micronecta) the Micronectidae as a separate taxon Siga- poweri and 8 males of M. (Micronecta) grise- rae. Subsequent authors did not recognize the ola – were collected in different water pools family or subfamily status of Micronectidae, e.g., Puton (1880), Oshanin (1912). Jaczewski and rivers in Bulgaria and fixed in 3:1 fixative (1924) introduced the modern name Micro- (96% ethanol - glacial acetic acid mixture). nectinae as a subfamily of Corixidae, and this The abdomens were dissected in 45% acetic status was used by nearly every author. Re- acid and the reproductive system structure was cently, Nieser (2002) raised the taxon to fami- analysed under stereoscope. Then the gonads ly status: Micronectidae. Over 150 species are were squashed in a small drop of 45% acetic described in the family worldwide; but over acid. The cover slips were removed by dry ice 100 of them belong to the genus Micronecta. technique. Slides were dehydrated in fresh According to Nieser, Chen (2006) the Mi- fixative (3:1) and air dried. To study the num- cronectidae consists of two subfamilies: the ber and the behaviour of the chromosomes the monotypic Synaptogobiinae (described from preparations were stained by Schiff-Giemsa Brazil) and Micronectinae with four genera method after Grozeva, Nokkala (1996). (Micronecta Kirkaldy, 1897 from the Old Chromosome spreads were analysed using World, Synaptonecta Lundblad, 1933 from a Laborlux 12 (Leitz, Wetzlar, Germany) mi- SE Asia and Florida, Monogobia Nieser et croscope with Olympus C 5060 digital cam- Chen, 2006, and Tenagobia Bergroth, 1899 era. from Tropical and Neotropical America). Ex- cept for two species, there are no cytogenetic RESULTS data on the Micronectidae. The chromosome formula of Micronecta poweri is reported by The male reproductive system of the three Southwood, Leston (1959) as 2n=24(22+XY); species was found to coincide with that de- Tenagobia (Fuscagobia) fuscata (Corixoidea, scribed and drawn for Nerthra terrestris (Ke- Micronectidae) has 2n=30(28+XY) and the van, 1948) (Nepomorpha: Gelastocoridae) by male meiosis is achiasmatic (Ituarte, Papes- Pendergrast (1957) as Mononyx: each testis chi, 2004). The information available on cy- comprises two lobes, their apical parts are totaxonomy of the family Micronectidae are snail-coiled and the long thick uncoiled strap- limited to these data. The spermatogenesis of like part crosses the abdomen to give rise to the the Micronecta species has not been studied vas deferens. It is an irregularly-coiled duct. earlier. Ducts from a pair of accessory glands enter In the present paper, the spermatogen- the apex of the long ductus ejaculatorius. esis of Micronecta (Dichaetonecta) scholtzi The number of chromosomes and their be- (Fieber, 1860), Micronecta (Micronecta) pow- haviour of all species are quite identical and eri (Douglas, Scott, 1869) and M. (Micronec- will be presented together. The adult males Comp. Cytogenet., 2008. 2(1) Achiasmatic male meiosis in Micronecta species 75 Figs 1-9. Male meiosis of Micronecta species. 1, 2 - early condensation stages of M. scholtzi. 3 - late conden- sation stage of M. scholtzi, showing 13 chromosome elements (11+XY). 4 - prometaphase I of M. scholtzi in side view. The bivalents consist of two side by side aligned chromosomes facing the opposite poles. 5 - metaphase I of M. griseola in side view. 6, 7 - second prometaphase (PMII) of M. scholtzi. The Y chromosome is clearly smaller than the X chromosome, and the sex chromosomes can be seen separately (6) or together (7). 8 - second metaphase (MII) of M. griseola, the sex chromosomes are quite similar in size, forming pseudobivalent in the middle of the plate. 9 - second anaphase (AII) of M. scholtzi, every anaphase group contains 12 chromosomes, eleven autosomes and a sex chromosome (X/Y). Bar=10μm. usually had the testes almost full of sperms consisted of side by side aligned homologous with a small number of dividing cells per chromosomes (Figs 1, 2), and the positively testis. No spermatogonial cells at metaphase heteropycnotic X and Y sex chromosomes con- were observed. Among the meiotic stages, nected to a nucleolus (Figs 1, 2). In some cells, post-pachytene stages were abundantly pres- in addition to sex chromosomes, 11 autosomal ent in the testes. Nuclei at early condensation bivalents could be traced (Fig. 2), hence hap- stages revealed autosomal bivalents, which loid chromosome number in these species is Comp. Cytogenet., 2008. 2(1) 76 S. Grozeva , N. Simov, S. Nokkala n=11+XY. Similar side by side association of alized or plesiomorphic condition in Heter- homologous chromosomes was still seen in bi- optera (Akingbohungbe, 1983). The analysis valents at late condensation stage, revealing 13 of reproductive system of Micronecta males chromosome elements (11+X+Y) (Fig. 3) and studied displayed a reduced follicle number prometaphase I (Fig. 4). In prometaphase, the as an apomorphy, and confirms the conclusion bivalents lay parallel to the equatorial plane, of Pendergrast (1957) that the reproductive or- the homologous chromosomes facing opposite gans of all aquatic bugs are of the same type. poles. This orientation of bivalents was still In Heteroptera, the existence of achias- observable in MI cells (Fig. 5). Clearly, chias- matic meiosis has been described in seven mata are absent and meiosis in males of these families, belonging to three infraorders: species is achiasmatic. At the second meta- Leptopodomorpha, in family Saldidae (Nok- phase (MII), autosomes did not form a clear kala, Nokkala, 1983); two superfamilies of ring but showed variable position from quite Cimicomorpha, Miroidea - families Miridae even distribution in prometaphase II (Figs 6, (Nokkala, Nokkala, 1986a) and Microphysi- 7) to a radial-like arrangement in MII (Fig. 8). dae (Nokkala, Grozeva, 2000); and Cimicoi- The sex chromosomes
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