I factors are responsible for the I-R system of hybrid dysgenesis in D
rosophila melanogaster. They belong to the LINE class of mobile elemen
ts, which transpose via reverse transcription of a full-length RNA int
ermediate. I factors are active members of the I element family, which
also contains defective I elements that are immobilized within peri-c
entromeric heterochromatin and represent very old components of the ge
nome. Active I factors have recently invaded natural populations of Dr
osophila melanogaster, giving rise to inducer strains. Reactive strain
s, devoid of active I factors, derive from old laboratory stocks estab
lished before the invasion. Transposition of I factors is activated at
very high frequencies in the germline of hybrid females issued from c
rosses between females from reactive strains and males from inducer st
rains. It results in the production of high rates of mutations and chr
omosomal rearrangements as well as in a particular syndrome of sterili
ty. The frequency of transposition of I factors is dependent on the am
ount of full-length RNA that is synthesized from an internal promoter.
This full-length RNA serves both as an intermediate of transposition
and presumably as a messenger for protein synthesis. Regulators of tra
nsposition apparently affect transcription initiation from the interna
l promoter. The data presented here lead to the proposal of a tentativ
e model for transposition.