RESUMO
Approximately 15% of eukaryotes contain supernumerary B chromosomes. When present, B chromosomes frequently represent as much as 5% of the genome. Despite thousands of reports describing the distribution of supernumeraries in various taxa, a comprehensive theory for the origin, maintenance, and evolution of B chromosomes has not emerged. Here, we sequence the complete genomes of individual cichlid fish (Astatotilapia latifasciata) with and without B chromosomes, as well as microdissected B chromosomes, to identify DNA sequences on the B. B sequences were further analyzed through quantitative polymerase chain reaction and in situ hybridization. We find that the B chromosome contains thousands of sequences duplicated from essentially every chromosome in the ancestral karyotype. Although most genes on the B chromosome are fragmented, a few are largely intact, and we detect evidence that at least three of them are transcriptionally active. We propose a model in which the B chromosome originated early in the evolutionary history of Lake Victoria cichlids from a small fragment of one autosome. DNA sequences originating from several autosomes, including protein-coding genes and transposable elements, subsequently inserted into this proto-B. We propose that intact B chromosome genes involved with microtubule organization, kinetochore structure, recombination and progression through the cell cycle may play a role in driving the transmission of the B chromosome. Furthermore, our work suggests that karyotyping is an essential step prior to genome sequencing to avoid problems in genome assembly and analytical biases created by the presence of high copy number sequences on the B chromosome.
Assuntos
Cromossomos/genética , Ciclídeos/genética , Proteínas de Peixes/genética , Genômica/métodos , Animais , Ciclídeos/classificação , Evolução Molecular , Duplicação Gênica , Sequenciamento de Nucleotídeos em Larga Escala , Cariótipo , Modelos Moleculares , Análise de Sequência de DNARESUMO
To better understand the genomic organization and evolution of Sox genes in vertebrates, we cytogenetically mapped Sox2 and Sox14 genes in cichlid fishes and performed comparative analyses of their orthologs in several vertebrate species. The genomic regions neighboring Sox2 and Sox14 have been conserved during vertebrate diversification. Although cichlids seem to have undergone high rates of genomic rearrangements, Sox2 and Sox14 are linked in the same chromosome in the Etroplinae Etroplus maculatus that represents the sister group of all remaining cichlids. However, these genes are located on different chromosomes in several species of the sister group Pseudocrenilabrinae. Similarly, the ancestral synteny of Sox2 and Sox14 has been maintained in several vertebrates, but this synteny has been broken independently in all major groups as a consequence of karyotype rearrangements that took place during the vertebrate evolution.
Assuntos
Ciclídeos/genética , Proteínas de Peixes/genética , Fatores de Transcrição SOXB1/genética , Fatores de Transcrição SOXB2/genética , Vertebrados/genética , Animais , Mapeamento Cromossômico , Proteínas de Peixes/classificação , Genoma/genética , Humanos , Hibridização in Situ Fluorescente , Filogenia , Especificidade da Espécie , Vertebrados/classificaçãoRESUMO
BACKGROUND: Cichlid fishes have been the subject of increasing scientific interest because of their rapid adaptive radiation which has led to an extensive ecological diversity and their enormous importance to tropical and subtropical aquaculture. To increase our understanding of chromosome evolution among cichlid species, karyotypes of one Asian, 22 African, and 30 South American cichlid species were investigated, and chromosomal data of the family was reviewed. RESULTS: Although there is extensive variation in the karyotypes of cichlid fishes (from 2n = 32 to 2n = 60 chromosomes), the modal chromosome number for South American species was 2n = 48 and the modal number for the African ones was 2n = 44. The only Asian species analyzed, Etroplus maculatus, was observed to have 46 chromosomes. The presence of one or two macro B chromosomes was detected in two African species. The cytogenetic mapping of 18S ribosomal RNA (18S rRNA) gene revealed a variable number of clusters among species varying from two to six. CONCLUSIONS: The karyotype diversification of cichlids seems to have occurred through several chromosomal rearrangements involving fissions, fusions and inversions. It was possible to identify karyotype markers for the subfamilies Pseudocrenilabrinae (African) and Cichlinae (American). The karyotype analyses did not clarify the phylogenetic relationship among the Cichlinae tribes. On the other hand, the two major groups of Pseudocrenilabrinae (tilapiine and haplochromine) were clearly discriminated based on the characteristics of their karyotypes. The cytogenetic mapping of 18S ribosomal RNA (18S rRNA) gene did not follow the chromosome diversification in the family. The dynamic evolution of the repeated units of rRNA genes generates patterns of chromosomal distribution that do not help follows the phylogenetic relationships among taxa. The presence of B chromosomes in cichlids is of particular interest because they may not be represented in the reference genome sequences currently being obtained.
Assuntos
Cromossomos , Ciclídeos/genética , Evolução Molecular , Animais , Mapeamento Cromossômico , Cariotipagem , RNA/genéticaRESUMO
Teleost fishes have become important models for studying the evolution of the genetic mechanisms of development. A key resource for comparative genomics and positional cloning are large-insert libraries constructed in bacterial artificial chromosomes. We have constructed bacterial artificial chromosome libraries for two species of teleost fish that are important models for the study of developmental evolution. Metriaclima zebra is one of several hundred closely related, morphologically diverse, haplochromine cichlids which have evolved over the last one million years in Lake Malawi, East Africa. The Mexican tetra, Astyanax mexicanus, is well known for adaptations related to the recent evolution of blind cave-dwelling forms. Clones and high-density filters for each library are available to the scientific community through the Hubbard Center for Genome Studies.