We have sequenced Bs), an insertion element isolated from a null allele of the Adhi locus encoding alcohol dehydrogenase in maize. The Bs) element is 3203 base pairs (bp) in length, has 302-bp identical long terminal direct repeats (LTRs), and created a 5-bp flanking direct duplication oftarget AdhI DNA upon insertion. The 5' LTR is followed by a canonical primer binding site with homology to the plant initiator methionyl-tRNA, and the 3' LTR is directly preceded by a polypurine stretch like that observed in retroviruses and retrotransposons. Bs) encodes two overlapping open reading frames specifying peptides of 740 and 168 amino acids. The longer open reading frame specifies a peptide with amino acid homology to the protease and nucleic acid binding moiety of retroviruses and retrotransposons. The deduced amino acid sequence encoded by Bsl lacks convincing homology to the polymerase (reverse transcriptase) encoded by retroposons, despite the fact that this polymerase-encoding domain is routinely the most conserved region of any such element. The sequence and relatively small size of Bs) suggest that this element is a deleted retrotransposon that inserted into AdhM with the aid of a reverse transcriptase function provided in trans. In vitro transcribed Bsl complementary RNA was translated in vitro to produce both a protein of 81 kDa representing open reading frame 1 (ORF1) and one of the 95-kDa size predicted for the frame-shifted fusion of ORF1 and ORF2. As with many other retroposons, the efficiency of translational initiation at the AUG beginning ORF1 was not noticeably affected by the presence of one or more upstream, unproductive AUGs in the complementary RNA transcript.By both structural and mechanistic criteria, eukaryotic insertion sequences and transposable elements can be divided into two major categories. One class of elements is bounded by inverted terminal repeat sequences and transposes through excision (1, 2) and/or enhanced replication (3) of the integrated element. Examples of this class of transposable elements are the controlling elements of maize (4) and the P elements of Drosophila (5). A second class of eukaryotic insertion sequences, the "retroposons," integrate into the chromosome after reverse transcription of element-encoded RNA (6), are particularly abundant in animals, and are the only type of insertion sequence yet discovered in fungi. Both the retroviruses (6) and the LI elements (7) of vertebrates and the "retrotransposons" Ty of yeast (8) and copia of Drosophila (9) fall into this latter category.Despite the early discovery and characterization of numerous distinct transposable elements in maize (1, 4), a canonical retrovirus or retrotransposon has not been described in plants. The DNA caulimoviruses ofplants have the structural properties and encode the reverse transcriptase activity definitional of a retrovirus or retrotransposon but apparently do not integrate into chromosomal DNA (10). Recently, Schwarz-Sommer et al. (11) have found that the Cin4 insertion element of m...