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The B Chromosome System of Omocestus Bolivari

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The B Chromosome System of Omocestus Bolivari Heredity 54 (1985) 385—390 The Genetical Society of Great Britain Received 14 January1985 TheB chromosome system of Omocestus bolivari: changes in B-behaviour in M4-polysomic B-males E. Viseras and Departamento de Genética, Facultad de Ciencias, J. P. M. Camacho Universidadde Granada, 18071 Granada, Spain. A metacentric B chromosome has been found in four out the five populations of Omocestus bolivari analysed. The B univalent frequently associates with the X chromosome at prophase I. The iso-B nature is deduced from the persistent association between the two arms of the B, but it is neither confirmed nor denied by its C-banding response. At first meiotic division B univalents sometimes divide equationally which lead to their loss in the form of microspermatids. These aspects of meiotic behaviour of the iso-B's are significantly influenced by the M4-polysomy. INTRODUCTION (AU, 2500 m, North side), 22 males at Alto del Chorrillo (ACH, 2700 m, South side) and 55 males Manyspecies of grasshopper carry supernumerary at La Alberquilla (LA, 2400 m, South side). (B) chromosomes as extra elements in their Testes were fixed in 1:3 acetic ethanol. In the cytogenetic systems (see Hewitt, 1979; Jones and CO population testes and gastric caeca of each Rees, 1982). These additional chromosomes are male were fixed following the method described usually heterochromatic, show positively by Kayano (1971). Females were injected with 0.05 heteropycnotic at first meiotic prophase, and often per cent colchicine in insect saline solution for six associate with the X chromosome leading in some hours before fixation of gastric caeca and ovarioles. instances to a preferential migration of both ele- All preparations for the routine cytogenetic analy- ments to the same anaphase I pole (Smith, 1953; sis of the material were made by squashing in 2 Henderson, 1961; Fontana and Vickery, 1973). per cent acetic orcein. Preparations for C-banding Omocestus bolivari is a grasshopper species were made and treated following the technique endemic in the Sierra Nevada (Granada, Spain) described in Camacho and Cabrero (1983). in which extra chromosomes have been detected in the form of autosome polysomy and iso-B chromosomes (Camacho, 1980; Camacho et a!., 1981). In the present report five natural popula- RESULTS tions of 0. bolivari are analysed for B chromosome distribution and frequency, meiotic behaviour of Themedium sized metacentric B chromosome of the iso-B's and their elimination in the form of 0. bolivari is an iso-chromosome, as was reported microspermatids. by Camacho eta!. (1981), on the basis of the similar length of its two arms and the meiotic pairing of them giving rise to ring B univalents (Camacho et MATERIAL AND METHODS a!., 1981). We have further investigated this matter using the C-banding technique. Unfortunately, the We caught 0. bolivari specimens in five well separ- B chromosome shows only a minute paracen- ated populations representative of the distribution tromeric C-band (fig. 1(a)). In consequence, since of this species: 40 males at Las Sabinas (LS, there are no interstitial C-bands which could serve 2100 m, North side of the Sierra Nevada), 18 males as chromosome markers, the C-banding pattern of and 19 females at Campos de Otero (CO, 2200 m, the B chromosome neither confirms nor denies its North side), 76 males at Albergue Universitario iso-chromosome nature. 386 E. VISERAS AND J. P. M. CAMACHO The iso-B chromosome was found in four out In some instances (like Myrmeleotettix macu- of the five natural populations analysed. So, the latus) B-frequency variation among populations is frequency of males carrying it ranged among popu- correlated with environmental characteristics lations from 0 in ACH to 28 per cent in CO (table (Hewitt and John, 1967). In, 0. ho/ivan, although 1). One out of 19 females analysed from CO carried a more detailed transect is necessary, as far as the one iso-B. In B-carrying males the iso-B chromo- data go a correlation seems to exist between B- some was found in both the somatic (gastric caeca, frequency and altitude. So, in the highest popula- fig. 1(e)) and germ lines (see fig. 1(a)), and its tion analysed (ACH, 2700 m) we have not found presence and number was constant in all cells in B chromosomes. The major B-frequency observed the carrier individuals which indicates it is a mitoti- is that of CO (2200 m), one of the lower popula- cally stable B chromosome. tions sampled. Intermediate altititudes have inter- mediate B-frequencies. The substantial difference, Table1 l-requencv of the iso-B in the populations besides temperature, between higher and lower populations is that in the former the snow covers Male types B-carrier Standard the ground for more time each year and, con- Population males males sequently, in them the period for grasshopper development, fertilisation and laying is shorter LS 3 750 37 than in lower populations. So, in higher popula- CO5 5 2778 13 AU 7 921 61 tions genotypes which more quickly complete their ACH 0 000 22 life-cycle will be favoured by natural selection. It LA 4 727 51 has been demonstrated that B chromosomes of the maculatus slow * One male carried 2B's grasshopper TYlyrineleotettix development (Harvey and Hewitt, 1979). Sig- Wehave analysed several aspects of meiotic nificantly, in our highest population (ACH) behaviour of the iso-B's in B-carrying males and individuals with extra chromosomes were not in two males from the AU population which were found, neither B chromosomes (this paper) nor moreover mosaic for M4-polysomy viz., non polysomic M4 autosomes (Viseras and Camacho, homologous associations between X, B and M4 1984), which indicates that zygotes carrying extra chromosomes may have reduced fitness in this (table 2), homologous association between the two environment. In lower populations there is snow arms of the iso-B (table 3),theformation of micro- for less time and any developmental retardation spermatids and the frequency of equational produced by extra chromosomes (B or M4) is less division of the B univalents during first meiotic crucial for survival of the individuals, as indicated division (table 4). Figs. I and 2 illustrate these by the presence in the AU population of two males aspects. carrying simultaneously B and extra M4 chromo- somes. DISCUSSION An interesting result is the influence of M4- polysomy on meiotic B-behaviour. Two B-carrying Todate, we have not detected any accumulation males from the AU population were also mosaic mechanism of the iso-B on the male side but, on for M4-polysomy, which is characterised by the the contrary, a B-elimination seems to occur during presence in some cells of heterochromatinised M4 spermatogenesis in the form of microspermatids. chromosomes (het-M4's). In these males the B Why the B chromosomes of 0. bolivariare takes part in heterochromatic non-homologous maintained in the populations despite their elimi- associations (with the X and the het-M4) at a lower nation we do not know. The existence of an frequency than in non polysomic B-males. Also accumulation mechanism in females similar to the two homologous arms of the iso-B pair much those described in Myrmeleotettix maculatus more frequently in M4-polysomic 13-males than in (Hewitt, 1973; 1976) and Me1anop1ustrnur-rubrum B-males. A similar correspondence between the (Lucov and Nur, 1973) might explain it. However, levels of homologous and non homologous associ- in the CO population we found about 28 per cent ations is apparent for het-M4 chromosomes. In this of B-carrying males and only 5percent of B- case, cells with 2 het-M4's show higher homologous carrying females, so that although an accumulation pairing along with lower non homologous associ- mechanism existed in females it would not he very ation than cells carrying a single het-M4. Another significant, given the low frequency of the B in change in meiotic B-behaviour in M4-polysomic them. B-males is their increased frequency of equational Table 2 X—B associationat first meiotic division in B-carrying males from AU population and in one male (AU 25) being furthermore M4-polysomic 0 % X-B 0/ X/B % Total X-B % X/B % Total I 0 B-carrying Zygotene 491 8768 69 1232 560 AU 25 males from Pachytene 427 7849 117 2151 544 Non-polysomic Diplotene 60 6000 40 4000 100 0 C') AU population Diplotene 37! 7541 12! 2459 492 follicles Metaphase I I 100 99 9900 100 0 Metaphase 1 10 196 509 9804 519 Total 1299 6142 816 3858 2115 Total 61 3050 159 6950 200 m ci) X-B-hM4 0/ X_B/hM4 % X-hM4/B % B-hM4/X 0/ X/B/hM4 0/ Total ci) -I Ill Follicles carrying Pachytene 7 3043 0 000 6 2609 3 1304 7 3043 23 I het-M Diplotene 11 2619 4 952 5 I 190 10 238! 12 2857 42 0 Metaphase I 0 000 0 000 6 600 0 000 94 9400 100 11 Total 18 109! 4 242 17 1030 13 788 113 6848 165 0 Follicles carrying Pachytene 2 714 0 000 3 1071 4 1429 19 6786 28 0 2 het-M4's Diplotene 3 789 2 526 3 789 4 1053 26 6842 38 0 1 0 000 0 000 2 870 0 000 2! ri Metaphase 9130 23 ci) Total 5 562 2 225 8 899 8 899 66 7416 89 —4 0) 0) 0 I— Table 3 Association between the two arms of the iso-B at first meiotic division in B-carrying males from AU population and in AU25 M4-polysomic B-carrying male Pachytene Diplotene Metaphase I Ring B- Straight B- % Ring B- Straight B- % Ring B- Straight B- % univalents univalents association univalents univalents association univalents univalents association B-carrying males from 331 40 8922 366 80 8206 246 274 4731 Au population AU 25 Non-polysomicfollicles 42 5 8936 88 12 8800 90 10 9000 Follicles carrying I het-M.
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