Evolution of the WANCY region in amniote mitochondrial DNA |
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Authors: | Seutin G; Lang BF; Mindell DP; Morais R |
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Institution: | Departement de Biochimie, Universite de Montreal, Quebec, Canada. |
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Abstract: | In most vertebrate mitochondrial genomes, the site for initiation of
light-strand replication, OL, is found within a cluster of five transfer
RNA (tRNA) genes (tRNA(Trp), tRNA(Ala), tRNA(Asn), tRNA(Cys), and
tRNA(Tyr)). This region and part of the adjacent cytochrome c oxydase
subunit I (COI) gene were sequenced for two crocodilian, two turtle, and
one snake species and for Sphenodon punctatus; part of the adjacent
nicotinamide adenine dinucleotide dehydrogenase subunit 2 (ND2) gene was
also sequenced for the crocodilian and turtle species. All had the typical
vertebrate gene order. The turtles and the snake have a lengthy noncoding
sequence between the tRNA(Asn) and tRNA(Cys) genes that we assumed to be
homologous to the mammalian OL. The crocodilians and Sphenodon lack such a
sequence, a condition they share with birds. Most proposed phylogenies for
the amniotes require that OL at this position was lost at least twice
during their diversification or was evolved independently more than once.
Within the five tRNA genes, frequencies of substitutions are much higher in
loops than in stems. Many loops vary dramatically in size among the
species; in the most extreme case, the D-arm of the Sphenodon tRNA(Cys) is
a "D-arm replacement" loop of seven nucleotides. Frequency of transitions
in stems is relatively uniform across tRNAs, but frequency of transversions
varies greatly. Mismatches in stems are infrequent, and their relative
frequency in a specific tRNA is unrelated to the frequency of substitution
in the corresponding gene. Several features of mammalian mitochondrial
tRNAs are conserved in WANCY tRNAs throughout amniotes. The inferred
initiation codon for COI is GTG in crocodilians, turtles, and the snake, a
condition they share with fishes, certain amphibians, and birds. TTG
appears to be the initiation codon for COI in Sphenodon; if correct, this
would be a novel initiation codon for vertebrate mitochondrial DNA.
Phylogenetic analyses of the inferred amino acid sequences of ND2 and COI
support the sister-group relationship of birds and crocodilians and suggest
that mammals are an early derived lineage within the amniotes.
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