Molecular Biology and Evolution, Vol 13, 261-277, Copyright © 1996 by Society for Molecular Biology and Evolution
EN Moriyama and JR Powell
We have summarized and analyzed all available nuclear DNA sequence
polymorphism studies for three species of Drosophila, D. melanogaster (24
loci), D. simulans (12 loci), and D. pseudoobscura (5 loci). Our major
findings are: (1) The average nucleotide heterozygosity ranges from about
0.4% to 2% depending upon species and function of the region, i.e., coding
or noncoding. (2) Compared to D. simulans and D. pseudoobscura (which are
about equally variable), D. melanogaster displays a low degree of DNA
polymorphism. (3) Noncoding introns and 3' and 5' flanking DNA shows less
polymorphism than silent sites within coding DNA. (4) X-linked genes are
less variable than autosomal genes. (5) Transition (Ts) and transversion
(Tv) polymorphisms are about equally frequent in non-coding DNA and at
fourfold degenerate sites in coding DNA while Ts polymorphisms outnumber Tv
polymorphisms by about 2:1 in total coding DNA. The increased Ts
polymorphism in coding regions is likely due to the structure of the
genetic code: silent changes are more often Ts's than are replacement
substitutions. (6) The proportion of replacement polymorphisms is
significantly higher in D. melanogaster than in D. simulans. (7) The level
of variation in coding DNA and the adjacent noncoding DNA is significantly
correlated indicating regional effects, most notably recombination. (8)
Surprisingly, the level of polymorphism at silent coding sites in D.
melanogaster is positively correlated with degree of codon usage bias. (9)
Three proposed tests of the neutral theory of DNA polymorphisms have been
performed on the data: Tajima's test, the HKA test, and the
McDonald-Kreitman test. About half of the loci fail to conform to the
expectations of neutral theory by one of the tests. We conclude that many
variables are affecting levels of DNA polymorphism in Drosophila, from
properties of nucleotides to population history and, perhaps, mating
structure. No simple, all encompassing explanation satisfactorily accounts
for the data.
ORIGINAL ARTICLE
Intraspecific nuclear DNA variation in Drosophila
Department of Biology, Yale University, New Haven, Connecticut 06520- 8104, USA.
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