3. OBJETIVOS ESPECÍFICOS
4.2.2.7. Análise das lâminas e idiogramas
As lâminas foram analisadas em fotomicroscópio de epifluorescência (OlympusTM BX51) com filtros apropriados para cada marcador, em aumento de 1.000x. As preparações foram capturadas pelo sistema digital Olympus DP73 com uso do software CellSens Standard 1.7 Full (Olympus Optical Co. Ltd.). Os cariótipos e idiogramas foram preparados utilizando o software Adobe Photoshop CS5 versão 12.0, seguindo as nomenclatura proposta por Levan et al. (1964).
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Capítulo 1
Unusual dispersion of histone repeats on the whole chromosomal
complement and their colocalization with ribosomal genes in Rachycentron
canadum (Rachycentridae, Perciformes)
Abstract Rachycentron canadum, the only representative of the family Rachycentridae, has been the focus of biotechnological interest due to its significant potential in marine fish farming. The chromosome set of this species has been widely investigated with respect to the location of genes and multigene families. A FISH analysis was performed using 4 multigene families as probes, represented by 5S and 18S ribosomal genes and histones H2B‐H2A and H3. Earlier data suggested that differential replication of heterochromatin could be partially associated with functional genes. Indeed, our results showed that the DNA contained in heterochromatic regions of R. canadum contains 5S and 18S ribosomal genes as well as the gene sequences of histones H2B‐H2A and H3, which were colocalized. The distribution of H3 sequences in all heterochromatic regions, except in 13q, could indicate an important evolutionary role for this class of repetitive sequences. Besides, the presence of chromosome regions bearing multifunctional repetitive sequences formed by H2B‐ H2A/H3/18S rDNA and H2B‐H2A/H3/5S rDNA clusters was demonstrated for the first time in fishes. The implications of differential histone gene extension and its functionality in the karyotype of R. canadum remain unknown.
Key words: Fish cytogenetics ∙ Heterochromatinization ∙ Karyotype evolution ∙ Multigene families ∙ Repetitive sequence expansion
Introduction
Rachycentron canadum (cobia), the only representative member of the monotypic
family Rachycentridae (Perciformes), is found in tropical seas worldwide. The high commercial value of this species, the facility of its spawning in captivity, its marked growth rate, good saline tolerance, and adaptability to confinement favor its intensive cultivation [Faulk and Holt, 2006; Liao and Leaño, 2007; Benetti et al., 2008]. Although farming methodology is constantly being improved, data available on the genome and the chromosomes of this species are still scarce [Jacobina et al., 2011].
Previous cytogenetic studies of R. canadum supplied information on karyotype structure, replication band patterns, and physical mapping of telomeric and ribosomal sequences [Jacobina et al., 2011]. However, the composition of heterochromatic portions of the genome and the possible presence of genes associated with these regions were hitherto unknown. Histones and ribosomal genes are vital multigene families in eukaryotes [Jordan et al., 2002; Roehrdanz et al., 2010]. Functionally, DNAhis (H1, H2A, H2B, H3, and H4) codes for a family of highly conserved, small basic proteins [Sellos et al., 1990; Chioda et al., 2002] that play an important role in the structural organization of chromatin which is essential in DNA packaging in the cell nucleus as well as in the regulation of gene expression. Physical mapping data of histone genes in fishes are largely unknown, with few studies focusing on this issue [Hashimoto et al., 2011; Lima‐Filho et al., 2012, 2014]. However, the multigene families of histones and ribosomal genes in some other organisms are known to exhibit integrated genome arrangements [Andrews et al., 1987; Roehrdanz et al., 2010]. Here, we investigated the chromosomal mapping of 5S and 18S ribosomal DNA (rDNA) sequences and histones H2B‐H2A and H3 in the chromosomes of R. canadum. Material and Methods Specimens, Chromosomal Preparations, and Banding
Forty specimens of R. canadum (Linnaeus, 1766) were used for cytogenetic analyses. The sample consisted of juveniles of undetermined sex, weighing approximately 35 g, obtained from a commercial fish farm in northeast Brazil.
There was no Animal Ethics Committee at the research institute; thus, this study was carried out in accordance with Brazilian law regarding the use of laboratory animals (Law No. 11.794/2008) and supported by the Brazilian Institute of Environment and Renewable Natural Resources (IBAMA, Process No. 556793/2009‐9). The authors underscore that the specimens were sacrificed under complete anesthesia (described in the next section), and all efforts were made to minimize suffering.
Specimens were previously submitted to in vivo mitotic stimulation for 24 h, using attenuated compound antigens [Molina, 2001; Molina et al., 2010]. After this period, the animals were anesthetized with clove oil (Eugenol), sacrificed by cervical dislocation followed by pithing. In detail, the fish were killed by inserting a thumb into their mouths, holding the fish with the opposite hand and displacing it dorsally. A metal spike was employed for pithing, pushed into the top of the head to destroy the brain and proximal end of the spinal cord. Death was recognized by cessation of respiration (opercular movement). Then the kidney tissue was removed. Metaphase chromosomes were obtained from cell suspensions of the anterior kidney by in vitro interruption of the mitotic cycle, in accordance with the methodology proposed by Gold et al. [1990]. A volume of 80 μl of cell suspension was dripped onto a slide covered with a film of distilled water heated to 60°C. Chromosomes