Both X chromosome inactivation and autosomal genomic imprinting generate a
functional hemizygosity. Here we consider models that explain the evolution
of genomic imprinting and X chromosome inactivation from novel perspective
s. Specifically, we suggest that random (in)activation events are common in
genes and gene clusters with a low probability of transcription. These gen
erate variability that natural selection has acted on to evolve stable mono
allelic expression. Possible selection forces might include a need for dosa
ge compensation and the prevention of biallelic silencing where a total swi
tch off would be lethal. Two different mechanisms can accomplish regular mo
noallelic expression - genomic imprinting and gene counting.