More than a decade ago, a surprising
observation was made in petunias. While trying to deepen the
purple color of these flowers, Rich Jorgensen and colleagues
introduced a pigment-producing gene under the control of a
powerful promoter. Instead of the expected deep purple color,
many of the flowers appeared variegated or even white. Jorgensen
named the observed phenomenon "cosuppression", since the expression
of both the introduced gene and the homologous endogenous gene
was suppressed (1-5).
First thought to be a quirk of petunias,
cosuppression has since been found to occur in many species
of plants. It has also been observed in fungi, and has been
particularly well characterized in Neurospora crassa,
where it is known as "quelling" (1-3).
But what causes this gene silencing effect?
Although transgene-induced silencing in some plants appears
to involve gene-specific methylation (transcriptional gene
silencing, or TGS), in others silencing occurs at the post-transcriptional
level (post-transcriptional gene silencing, or PTGS). Nuclear
run-on experiments in the latter case show that the homologous
transcript is made, but that it is rapidly degraded in the
cytoplasm and does not accumulate (1, 3, 6).
Introduction of transgenes can trigger
PTGS, however silencing can also be induced by the introduction
of certain viruses (2, 3).
Once triggered, PTGS is mediated by a diffusible, trans-acting
molecule. This was first demonstrated in Neurospora,
when Cogoni and colleagues showed that gene silencing could
be transferred between nuclei in heterokaryotic strains (1, 7).
It was later confirmed in plants when Palauqui and colleagues
induced PTGS in a host plant by grafting a silenced, transgene-containing
source plant to an unsilenced host (8).
From work done in nematodes and flies, we now know that the
trans-acting factor responsible for PTGS in plants is dsRNA
(1-3).
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