The Fish Specific Genome Duplications and Fish Diversity
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Previous studies suggested that there were two rounds of genome duplication early in vertebrate history(the 2R hypothesis).And recent phylogenetic and comparative data suggested that an additional whole genome duplication occurred in the stem lineage of ray-finned(actinopterygian) fishes,but not in that of the land vertebrates,namely the fish-specific genome duplication(the 3R hypothesis).The genome duplication issues are intriguing in organism evolutionary research currently.And the role of genome duplication in evolution are still controversial.The teleost fish is composed of more than half of extant vertebrate species and displays remarkable variation in morphological and physiological adaptations.The genomic complexity of the teleosts might be the reason for their evolutionary success and astonishing biological diversity.Keywords:
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Comparative Genomics
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Subfunctionalization
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The presence of additional hox clusters in the zebrafish has led to the hypothesis that there was a whole genome duplication at the origin of modern fish. To investigate the generality of this assumption, we analyzed all available actinopterygian fish gene families, and sequenced nuclear receptors from diverse teleost fish. The origin and timing of duplications was systematically determined by phylogenetic analysis. More genes are indeed found in zebrafish than in mouse. This abundance is shared by all major groups of euteleost fish, but not by eels. Phylogenetic analysis shows that it may result from frequent independent duplications, rather than from an ancestral genome duplication. We predict two zebrafish paralogs for most mouse or human genes, thus expressing a note of caution in functional comparison of fish and mammalian genomes. Redundancy appears to be the rule in fish developmental genetics. Finally, our results imply that the outcome of genome projects cannot be extrapolated easily between fish species.
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The connection between increasing genomic complexity by gene duplication and origin of vertebrates is of great interest to evolutionary biologists. The most widely accepted hypothesis is that the origin of vertebrates correlated with two rounds of genome duplication early in vertebrate evolution, which is originally suggested by Ohno in 1970 and later modified by Holland et al in 1994. Three important observations have been marshaled in line with this hypothesis. First, the number of coding genes in vertebrates is roughly four times the number estimated for invertebrates such as Drosophila and ascidian. Moreover, for many genes in the invertebrates such as Drosophila and amphioxus, there are typically four putative vertebrate orthologs. Second, there exists one Hox gene cluster in invertebrates such as arthropods and echinoderms and cephochordates such as amphioxus and four clusters in vertebrates, although teleost fishes are exceptional with the recent discovery of seven Hox gene clusters. Third, large sections of chromosomes have similar gene order between fishes and mammals, and there is evidence of paralogy of chromosomes in mammalian genomes. It is proposed that the first round of gene duplication occurred after the diversification of vertebrates from cephalochordate, and the second round after the diversification of jawed vertebrates from jawless vertebrates. However, it remains open whether gene duplication early in vertebrate evolution is small-scale (regional) DNA duplication or large-scale (genomic) duplication.
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It is now clear that a whole-genome duplication (WGD) occurred at the base of the teleost fish lineage. Like the other anciently polyploid genomes investigated so far, teleost genomes now behave like diploids with chromosomes forming pairs at meiosis. The diploidization process is currently poorly understood. It is associated with many gene deletions, such that one of the duplicates is lost at most loci and has also been proposed to coincide with an increase in genomic instability. Here we ask whether WGD is a determinant of the genomic rearrangement rate in teleosts. We study variability of the rates of rearrangement along a vertebrate phylogenetic tree, composed of 3 tetrapods (human, chicken, and mouse) and 3 teleost fishes (zebrafish, Tetraodon, and Takifugu), whose complete genome sequences are available. We devise a simple parsimony method for counting rearrangements, which takes into account various methodological complications caused by the WGD and the subsequent gene losses. We show that there does appear to be an increase in rearrangement rate after WGD, but that there is also a great deal of additional variability in rearrangement rates across species.
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Abstract An important mechanism for the evolution of phenotypic complexity, diversity and innovation, and the origin of novel gene functions is the duplication of genes and entire genomes. Recent phylogenomic studies suggest that, during the evolution of vertebrates, the entire genome was duplicated in two rounds (2R) of duplication. Later, ∼350 mya, in the stem lineage of ray‐finned (actinopterygian) fishes, but not in that of the land vertebrates, a third genome duplication occurred—the fish‐specific genome duplication (FSGD or 3R), leading, at least initially, to up to eight copies of the ancestral deuterostome genome. Therefore, the sarcopterygian (lobe‐finned fishes and tetrapods) genome possessed originally only half as many genes compared to the derived fishes, just like the most‐basal and species‐poor lineages of extant fishes that diverged from the fish stem lineage before the 3R duplication. Most duplicated genes were secondarily lost, yet some evolved new functions. The genomic complexity of the teleosts might be the reason for their evolutionary success and astounding biological diversity. BioEssays 27:937–945, 2005. © 2005 Wiley Periodicals, Inc.
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Multiple Chromosomal Rearrangements Structured the Ancestral Vertebrate Hox-Bearing Protochromosomes
While the proposal that large-scale genome expansions occurred early in vertebrate evolution is widely accepted, the exact mechanisms of the expansion--such as a single or multiple rounds of whole genome duplication, bloc chromosome duplications, large-scale individual gene duplications, or some combination of these--is unclear. Gene families with a single invertebrate member but four vertebrate members, such as the Hox clusters, provided early support for Ohno's hypothesis that two rounds of genome duplication (the 2R-model) occurred in the stem lineage of extant vertebrates. However, despite extensive study, the duplication history of the Hox clusters has remained unclear, calling into question its usefulness in resolving the role of large-scale gene or genome duplications in early vertebrates. Here, we present a phylogenetic analysis of the vertebrate Hox clusters and several linked genes (the Hox "paralogon") and show that different phylogenies are obtained for Dlx and Col genes than for Hox and ErbB genes. We show that these results are robust to errors in phylogenetic inference and suggest that these competing phylogenies can be resolved if two chromosomal crossover events occurred in the ancestral vertebrate. These results resolve conflicting data on the order of Hox gene duplications and the role of genome duplication in vertebrate evolution and suggest that a period of genome reorganization occurred after genome duplications in early vertebrates.
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Rhodopsin mediates an essential step in image capture and is tightly associated with visual adaptations of aquatic organisms, especially species that live in dim light environments (e.g., the deep sea). The rh1 gene encoding rhodopsin was formerly considered a single-copy gene in genomes of vertebrates, but increasing exceptional cases have been found in teleost fish species. The main objective of this study was to determine to what extent the visual adaptation of teleosts might have been shaped by the duplication and loss of rh1 genes. For that purpose, homologous rh1/rh1-like sequences in genomes of ray-finned fishes from a wide taxonomic range were explored using a PCR-based method, data mining of public genetic/genomic databases, and subsequent phylogenomic analyses of the retrieved sequences. We show that a second copy of the fish-specific intron-less rh1 is present in the genomes of most anguillids (Elopomorpha), Hiodon alosoides (Osteoglossomorpha), and several clupeocephalan lineages. The phylogenetic analysis and comparisons of alternative scenarios for putative events of gene duplication and loss suggested that fish rh1 was likely duplicated twice during the early evolutionary history of teleosts, with one event coinciding with the hypothesized fish-specific genome duplication and the other in the common ancestor of the Clupeocephala. After these gene duplication events, duplicated genes were maintained in several teleost lineages, whereas some were secondarily lost in specific lineages. Alternative evolutionary schemes of rh1 and comparison with previous studies of gene evolution are also reviewed.
Human evolutionary genetics
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Segmental duplication
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