Are macronuclear subunits in Paramecium functionally diploid?

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Abstract
SUMMARY: The organization of the genetic material in the macronucleus of ciliates has been the subject of considerable controversy. Two of the four models of macronuclear structure predict assortment of the alternative phenotypes of heterozygotes during vegetative growth. Early studies of the phenotypic behaviour of heterozygotes after macronuclear regeneration (Sonneborn, 1947) had supported a diploid subunit model. The availability of quantitative predictions of the rate of assortment for the haploid and chromosomal models (Preer, this volume) and the existence of two alleles controlling a quantitative trait, copper tolerance, in Paramecium tetraurelia, has provided an opportunity to test these models. The median tolerance limits to copper of unselected sublines were measured as a function of age. There was no increase in the variance among sublines, as the haploid and chromosomal models predict. Quantitative evaluation shows that subdiploid models with a kinetic complexity of 860 or less are not compatible with the results. This experiment was not sensitive enough, however, to exclude subdiploid models if the kinetic complexity is 2000 or greater. Selection on heterozygotes also failed to provide evidence in favour of assortment. All the results are consistent with and support the diploid subunit model of the Paramecium macronucleus.