Complete nucleotide sequence of the mitochondrial genome of a salamander, Mertensiella luschani
Introduction
Salamanders, the tailed amphibians (order Caudata), may represent a living paradigm of the tempo and mode in which the adaptations that are required to live on land were acquired in the evolution of vertebrates. Many of the 400 or so living salamander species (Frost, 1985) display peculiar feeding, breathing, urogenital, and reproductive apparati, which can be interpreted as alternative or sequential morphological adaptations to terrestrial life that were evolved when sarcopterygian ancestors of amphibians colonized land in the Devonian (360 MYA). Moreover, salamanders are also characterized by a wide variety of gametogenetic, mating, developmental, and parental care strategies (Duellman and Trueb, 1994), which can be perceived as an adaptation to permit reproduction on terrestrial conditions.
To trace and better understand the sequence of evolutionary events that produced the diversity in life modes exhibited by salamanders, it is mandatory first to infer the phylogenetic relationships of the group with other living amphibians (i.e. frogs and caecilians) and with other vertebrates (Titus and Larson, 1995). Therefore, we determined the complete sequence of the mitochondrial genome of an old world salamander, Mertensiella luschani. The usefulness of these new mitochondrial sequence data in resolving controversies on amphibian phylogenetic relationships was addressed elsewhere (Zardoya and Meyer, 2001). Here, we present a formal description of the molecular and structural features of the salamander mitochondrial genome and compared it with the complete mitochondrial genomes of other relevant jawed vertebrates.
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Mitochondrial DNA extraction
Mitochondrial DNA (mtDNA) was purified from eggs and liver of six specimens of the salamander species M. luschani, as previously described (Zardoya et al., 1995). After homogenization, mitochondria were separated from intact nuclei and cellular debris by a low-speed centrifugation. Mitochondrial DNA was extracted from isolated mitochondria by a standard alkaline lysis procedure, and cleaved with EcoRI and HindIII restriction enzymes. Two EcoRI fragments of 1.3 (comprising the complete control
Organization of the salamander mitochondrial genome
The mitochondrial genome of the salamander M. luschani is 16,650 bp long and conforms to the consensus vertebrate mitochondrial gene order (Fig. 1). As in other vertebrates, 2 rRNAs, 22 tRNAs, and 13 proteins are encoded by the salamander mitochondrial genome (Fig. 1). The overall base composition of the L-strand is A: 32%, T: 29%, C: 24%, and G: 15%. There are only nine noncoding intergenic spacer nucleotides. Interestingly, this paucity of intergenic spacers contrasts with the existence of a
Acknowledgments
Meredith J. Mahoney gave very useful advice on primers and PCR conditions to amplify the salamander ND4 gene. We thank two anonymous reviewers for their useful comments on an earlier version of the paper. R.Z. was sponsored by a postdoctoral contract of the Ministerio de Educacion y Cultura of Spain. This work received partial financial support from grants from the Lion Foundation, the Deutsche Forschungs Gemeinschaft (ME 1725/4-1), the University of Konstanz, the US National Science Foundation
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