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AN APPROACH TO POPULATION AND EVOLUTIONARY GENETIC THEORY FOR GENES IN MITOCHONDRIA AND CHLOROPLASTS, AND SOME RESULTS
C. William Birky Jr. 1, Takeo Maruyama 2, and Paul Fuerst 1
1 Department of Genetics, The Ohio State University, Columbus,
Ohio 43210
2 National Institute of Genetics, Mishima 411, Japan
We developed population genetic theory for organelle genes, using
an infinite alleles model appropriate for molecular genetic data, and considering
the effects of mutation and random drift on the frequencies of selectively
neutral alleles. The effects of maternal inheritance and vegetative segregation
of organelle genes are dealt with by defining new effective gene numbers,
and substituting these for 2Ne in classical theory of
nuclear genes for diploid organisms. We define three different effective gene
numbers. The most general is N
, defined as a function
of population size, number of organelle genomes per cell, and proportions
of genes contributed by male and female gametes to the zygote. In many organisms,
vegetative segregation of organelle genomes and intracellular random drift
of organelle gene frequencies combine to produce a predominance of homoplasmic
cells within individuals in the population. Then, the effective number of
organelle genes is Neo, a simple function of the numbers
of males and females and of the maternal and paternal contributions to the
zygote. Finally, when the paternal contribution is very small, N
eo is closely approximated by the number of females, N
f. Then if the sex ratio is 1, the mean time to fixation or loss
of new mutations is approximately two times longer for nuclear genes than
for organelle genes, and gene diversity is approximately four times greater.
The difference between nuclear and organelle genes disappears or is reversed
in animals in which males have large harems. The differences between nuclear
and organelle gene behavior caused by maternal inheritance and vegetative
segregation are generally small and may be overshadowed by differences in
mutation rates to neutral alleles. For monoecious organisms, the effective
number of organelle genes is approximately equal to the total population size
N. We also show that a population can be effectively subdivided for organelle
genes at migration rates which result in panmixis for nuclear genes, especially
if males migrate more than females.
Accepted on November 1, 1982
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