Ea. Kellogg et R. Appels, INTRASPECIFIC AND INTERSPECIFIC VARIATION IN 5S RNA GENES ARE DECOUPLED IN DIPLOID WHEAT RELATIVES, Genetics, 140(1), 1995, pp. 325-343
5S RNAs form part of the ribosome in most organisms. In some, e.g., pr
okaryotes and some fungi, the genes are part of the ribosomal operon,
but in most eukaryotes they are in tandem arrays of hundreds to thousa
nds of copies separate from the main ribosomal array. 5S RNA genes can
be aligned across kingdoms. We were therefore surprised to find that,
for 28 diploid species of the wheat tribe (Triticeae), nucleotide div
ersity within an array is up to 6.2% in the genes, not significantly d
ifferent from that of the nontranscribed spacers. Rates of concerted e
volution must therefore be insufficient to homogenize the entire array
. Between species, there are significantly fewer fixed differences in
the gene than would be expected, given the high within-species variati
on. In contrast, the amount of variation between species in the spacer
is the same as or greater than that within individuals. This leads to
a paradox. High variation within an individual suggests that there is
little selection on any particular gene within an array. But conserva
tion of the gene across species implies that polymorphisms are periodi
cally eliminated at a rate approximately equal to or greater than that
of speciation. Levels of intraspecific polymorphism and interspecific
divergence are thus decoupled. This implies that selective mechanisms
exist to eliminate mutations in the gene without also affecting the s
pacer.