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Cytogenetic Report on Cordulegaster Brevistigma and Watanabeopetalia Atkinsoni1st June 2019101

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Cytogenetic Report on Cordulegaster Brevistigma and Watanabeopetalia Atkinsoni1st June 2019101 Cytogenetic report on Cordulegaster brevistigma and Watanabeopetalia atkinsoni1st June 2019101 Cytogenetic report onCordulegaster brevistigma and Watanabeopetalia atkinsoni (Odonata: Cordulegastridae, Chlorogomphidae) Gurinder Kaur Walia1 & Sarabjit Singh Chahal Department of Zoology and Environmental Sciences, Punjabi University, Patiala 147002, Punjab, India; 1 corresponding author, [email protected] Received 26th October 2018; revised and accepted 8th April 2019 Abstract. Live adult male specimens of C. brevistigma and W. atkinsoni have been collected from Shimla, Himachal Pradesh (India). Male germ cell chromosomes of the species are de- scribed on the basis of conventional staining, C-banding, silver nitrate staining and sequence specific staining. Both the species possesses 2n = 25m, as a diploid chromosome number and XO (♂)/XX (♀) type sex determination. In both the species, dark terminal C-bands are present on all the autosomal bivalents and X chromosome is C-positive throughout the length. Terminal light/dark NORs (Nucleolar Organizer Regions) are present on all auto- somal bivalents, while X chromosome also possesses terminal NORs. During sequence spe- cific staining, all the autosomal bivalents show prominent terminal DAPI (4',6-diamidino- 2-phenylindole) and CMA3 (Chromomycin A3) bright regions and X chromosome also pos- sesses both DAPI and CMA3 signals. In addition, a brief review of the size of X element in the allied families Cordulegastridae, Corduliidae and Macromiidae is given. Further key words. Dragonfly, Anisoptera, chromosomes, conventional staining, C-band- ing, silver nitrate staining, sequence specific staining, India Introduction Cytogenetic data on eight species of the Cordulegastridae is available world- wide (Oguma 1930; Oksala 1939a, b; Kichijo 1942b; Kiauta 1968a, b, 1969; Cruden 1968, 1969; Kiauta & Kiauta 1976; Zhu & Wu 1986; Sandhu & Malhotra 1994; Perepelov et al. 2001), whereas no report is available for the Chlorogomphidae. In the present study, Cordulegaster brevi stigma Selys, 1854, (Cordulegastridae) and Watanabeopetalia atkinsoni (Selys, 1878), (Chlorogomphidae) have been studied cytogentically. Struc- ture and behaviour of chromosomes during meiosis, detection of constitu- tive heterochromatin, localization of NORs (Nucleolar Organizer Regions) Odonatologica 48(1/2) 2019: 101-113Odonatologica – DOI:10.5281/zenodo.2677693 48(1/2) 2019: 101-113 102 G. Kaur Walia & S. Singh Chahal and distribution of AT- and GC-rich regions were investigated and com- pared in both species. Material and methods Live adult male specimens of Cordulegaster brevistigma and Watanabeopeta­ lia atkinsoni were collected from Summer Hills, Shimla, Himachal Pradesh, India, in June 2015. Specimens were dissected in 0.67 % saline solution in the field and testes were taken out. Subsequently, testes were put in sodium citrate (0.9 %) for 45 minutes and then fixed in freshly prepared Carnoy’s fixative (3:1, absolute alcohol: glacial acetic acid). Testes were teased on grease free slides and slides were air dried. Slides were prepared for con- ventional staining (Carr & Walker 1961), C-banding (Sumner 1972), sil- ver nitrate staining (Howell & Black 1980) and sequence specific staining (Rebagaliati et al. 2003). Relevant meiotic and mitotic stages were micro- photographed. Results Conventional staining In both studied species, 25 elements were present during spermatogonial metaphase; out of these, 24 were autosomes, also including a pair of m chro- mosomes and one X chromosome. X chromosome was the 2nd smallest ele- ment in Cordulegaster brevistigma (Fig. 1a), while it was the largest element in the whole complement in Watanabeopetalia atkinsoni (Fig. 1e). Dur- ing the diplotene stage of W. atkinsoni autosomal bivalents showed cross- shaped structures due to the presence of single chiasma per bivalent, while a large bipartite X chromosome and small m bivalent were clearly visible (Fig. 1f). In both species, during diakinesis and metaphase I, 13 elements were visible; among these, 12 were autosomal bivalents and one was an X chromosome (Figs 1b, c, g, h). During metaphase II in C. brevistigma the chromosomes were half the size of metaphase I chromosomes, while X and m chromosomes were clearly distinguishable (Fig. 1d). Banding patterns In C. brevistigma, during diplotene and diakinesis, all the autosomal biva- lents including m bivalent showed terminal C-bands and the X chromosome Odonatologica 48(1/2) 2019: 101-113 Cytogenetic report on Cordulegaster brevistigma and Watanabeopetalia atkinsoni 103 was C-positive and showed bipartite behaviour in the diplotene (Figs 2a, b). In W. atkinsoni, in metaphase I, all the autosomal bivalents showed termi- nal C-bands, while m bivalent and X chromosome were C-positive because of condensation of elements (Figs 2c, d). In C. brevistigma, during diplo- tene, terminal NORs were present on all the autosomal bivalents and the X chromosome showed light NOR bands on both the chromatids (Fig. 2e). During diakinesis, all the autosomal bivalents showed terminal NORs and the X chromosome was rich in NORs (Fig. 2f). In W. atkinsoni, during dia- kinesis, terminal NORs were present on autosomal bivalents and the large X chromosome showed dark NOR bands (Figs 2g, h). In both the species, the interphase cell showed DAPI and CMA3 bright regions (Figs 3a, b, e, f) and all the autosomal bivalents showed terminal DAPI (4',6-diamidino-2- phenylindole) and CMA3 (Chromomycin A3) bright regions and the X chro- mosome was also DAPI and CMA3 bright in diakinesis (Figs 3c, d, g, h). Discussion Cytogenetically, Odonata are characterized by holokinetic chromosomes, post-reductional meiosis for sex chromosome, XX/XO (Female/Male) sex determining mechanism, with m chromosomes present and a single chi- asma per bivalent. So far, only eight species of Cordulegastridae have been cytogenetically studied world-wide (Table 1). The majority of the species possesses the chromosome number 2n(♂) = 25 (Oguma 1930; Oksala 1939a, b; Kichijo 1942b; Kiauta 1968a, b, 1969; Cruden 1968, 1969; Ki- auta & Kiauta 1976; Zhu & Wu 1986; Perepelov et al. 2001). The only exception is Anotogaster basalis with a chromosome number 2n = 23, which originates by the fusion of m chromosomes with another pair of autosomes (Sandhu & Malhotra 1994). Among these, four species of Cordulegaster possess 2n = 25m, which is the type number of the family. In the present study, C. brevistigma also possesses 2n (♂) = 25m, with X0(♂)/XX(♀) type sex determination, which is in accordance with reports of the chromosome number of other species in the genus. X is the 2nd smallest element in the complement after m chromosomes. No cytogenetic report was hitherto available on any species of the Chlorogomphidae. This study shows Watan­ abeopetalia atkinsoni possesses 2n (♂) = 25m, with X0(♂)/XX(♀) sex deter- mination and X is the largest element in the complement. Odonatologica 48(1/2) 2019: 101-113 104 G. Kaur Walia & S. Singh Chahal Figure 1. a−d, Conventional staining in Cordulegaster brevistigma; e−h, conven- tional staining in Watanabeopetalia atkinsoni. a − Spermatogonial metaphase; b, c – diakinesis; d − metaphase I; e − spermatogonial metaphase; f – diplotene; g, h − diakinesis. X and m marked with arrows. Bar: 0.01 mm Odonatologica 48(1/2) 2019: 101-113 Cytogenetic report on Cordulegaster brevistigma and Watanabeopetalia atkinsoni 105 Figure 2. a−b, C-banding in Cordulegaster brevistigma; c−d, C-banding in Watana­ beopetalia atkinsoni; e−f, silver nitrate staining in Cordulegaster brevistigma; g−h, silver nitrate staining in Watanabeopetalia atkinsoni. a – Diplotene; b – diakinesis; c, d − metaphase I; e – diplotene; f, g, h − diakinesis. X and m marked with arrows. Bar: 0.01 mm Odonatologica 48(1/2) 2019: 101-113 106 G. Kaur Walia & S. Singh Chahal Figure 3. a−d, Sequence specific staining in Cordulegaster brevistigma; e−h, se- quence specific staining in Watanabeopetalia atkinsoni. a, c, e, g − DAPI staining; b, d, f, h − CMA3 staining; a, b, e, f − interphase cell; c, d, g, h − diakinesis. X and m marked with arrows. Bar: 0.01 mm Odonatologica 48(1/2) 2019: 101-113 Cytogenetic report on Cordulegaster brevistigma and Watanabeopetalia atkinsoni 107 Size of the X chromosome in Cordulegastridae, Corduliidae and Macro- miidae In the Cordulegastridae, the size of X chromosome has been estimated in six out of eight cytogentically studied species. It is smallest when m chro- mosomes are absent (Kiauta & Kiauta 1976; Zhu & Wu 1986; Sandhu & Malhotra 1994) and 2nd smallest when m chromosomes are present (Oguma 1930; Cruden 1968, 1969). The only exception is Anotogaster sie­ boldii, in which X chromosome shows variation in size, as it is 2nd smallest (Oguma 1930) and medium sized (Perepelov et al. 2001). In the Cordulii- dae, the size of the X chromosome has been reported only in 15 out of 23 cy- togenetically studied species. It is smallest or 2nd smallest in majority of spe- cies. The only exception is Somatochlora borisi with chromosome comple- ment (n(♂) = 11/neo-XY) which originated from two fusions, one between two small pairs of autosomes and another between a pair of autosomes and the X element (Grozeva & Marinov 2007). In family Macromiidae, among the four cytogenetically studied species, the X chromosome is the 2nd small- est in three species (Dasgupta 1957; Cruden 1968; Kiauta 1977; Walia & Chahal 2018). The size of the X chromosome has been reviewed in the three allied fami- lies Cordulegastridae, Corduliidae and Macromiidae (Table
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