David M. Rand, Michele Dorfsman and Lisa M. Kann
Department of Ecology and Evolutionary Biology
Brown University
Providence, RI 02912
Address for correspondence:
Dr. David M. Rand
Department of Ecology and Evolutionary Biology
Brown University
Box G-W
Providence, RI 02912
Phone (401) 863-2890
Fax (401) 863-2166
email David_Rand@brown.edu
Key words: mtDNA / neutral theory / Drosophila / selection / nucleotide
polymorphism / molecular evolution
Running Head: Neutrality tests of mtDNA
Abstract
To test hypotheses of neutral evolution of mitochondrial DNA
(mtDNA), nucleotide sequences were determined for 1515 base pairs of the
NADH dehydrogenase subunit 5 (ND5) gene in the mitochondrial DNA of 29
lines of Drosophila melanogaster and nine lines of its sibling species
D. simulans. In contrast to the patterns for nuclear genes, where
D. melanogaster generally exhibits much less nucleotide polymorphism, the
number of segregating sites was slightly higher in a global sample of nine
ND5 sequences in D. melanogaster (s = 8) than in the nine lines of D. simulans
(s = 6). When compared to variation at nuclear loci, the mtDNA variation
in D. melanogaster does not depart from neutral expectations. The
ND5 sequences in D. simulans, however, show fewer than half the number
of variable sites expected under neutrality when compared to sequences
from the period locus. While this reduction in variation is not significant
at the 5% level, HKA tests with published restriction data for mtDNA
in D. simulans do show a significant reduction of variation suggesting
a selective sweep of variation in the mtDNA in this species. Tests
of neutral evolution based on the ratios of synonymous and replacement
polymorphism and divergence are generally consistent with neutral expectations,
although a significant excess of amino acid polymorphism within both species
is localized in one region of the protein. The rate of mtDNA evolution
has been faster in D. melanogaster than in D. simulans and the population
structure of mtDNA is distinct in these species. The data reveal
how different rates of mtDNA evolution between species and different histories
of neutral and adaptive evolution within species can compromise historical
inferences in population and evolutionary biology.