Rb. Flavell, INACTIVATION OF GENE-EXPRESSION IN PLANTS AS A CONSEQUENCE OF SPECIFIC SEQUENCE DUPLICATION, Proceedings of the National Academy of Sciences of the United Statesof America, 91(9), 1994, pp. 3490-3496
Numerous examples now exist in plants where the insertion of multiple
copies of a transgene leads to loss of expression of some or all copie
s of the transgene. Where the transgene contains sequences homologous
to an endogenous gene, expression of both transgene and endogenous gen
e is sometimes found to be impaired. Several examples of these phenome
na displaying different features are reviewed. Possible explanations f
or the observed phenomena are outlined, drawing on known cellular proc
esses in Drosophila, fungi, and mammals as well as plants. It is hypot
hesized that duplicated sequences can, under certain circumstances, be
come involved in cycles of hybrid chromatin formation or other process
es that generate the potential for modification of inherited chromatin
structure and cytosine methylation patterns. These epigenetic changes
could lead to altered transcription rates or altered efficiencies of
mRNA maturation and export from the nucleus. Where the loss of gene ex
pression is posttranscriptional, antisense RNA could be formed on accu
mulated, inefficiently processed RNAs by an RNA-dependent RNA polymera
se or from a chromosomal promoter and cause the observed loss of homol
ogous mRNAs and possibly the modification of homologous genes. It is s
uggested that the mechanisms evolved to help silence the many copies o
f transposable elements in plants. Multicopy genes that are part of th
e normal gene catalog of a plant species must have evolved to avoid th
ese silencing mechanisms or their consequences.