Six independent derivatives of the bz-ml3 allele, which contains a 2.2-kilobase-pair defective Suppressormutator (dSpm) insertion at the bronze (bz) locus, have been isolated and analyzed. The derivatives were selected for alterations in the frequency and timing of somatic reversion; such derivatives have previously been analyzed genetically and designated "changes in state" by McClintock [McClintock, B. (1955) Carnegie Inst. Washington, Yearb. 54, 245-255]. All of the derivatives analyzed in the present study revert substantially later in development than the original insertion mutation and some show a very low frequency of reversion as well. All of the derivatives contain insertions at the same site as the parent bz-m13 allele. Deletions of 400-1300 base pairs were found in the dSpm elements in four of the six derivatives; the remaining derivatives could not be distinguished structurally from the original mutant allele. The results suggest that changes in the frequency and developmental timing of excision are attributable to alterations in the dSpm element. Furthermore, these data suggest that DNA sequences near the ends of the element are important for responding to the two transacting functions supplied by the transposition-competent Suppressor-mutator (Spm) element.Transposable elements were first identified and described in maize (1, 2). Early work revealed the presence of distinct families of elements, each composed of members able to transpose on their own (transposition-competent elements) and members unable to transpose in the absence of a transposition-competent member (defective elements; for reviews see refs. 3 and 4). Recent molecular analyses of maize transposable elements have shown that the defective elements of a family are often deletion derivatives of the transposition-competent elements (5-7).Mutations caused by the insertion of transposition-competent elements in or near a locus are unstable, reverting both somatically and germinally at a high frequency. Mutations caused by insertions of defective elements are unstable only when a transposition-competent element of the same family is present in the genome. A characteristic of each unstable mutation is its particular pattern of somatic reversion. McClintock described the isolation of derivatives of insertion mutations that showed heritable changes in the frequency and developmental timing of somatic reversion. She designated the new unstable alleles that arose in this manner "changes in state" (8, 9). Since then, derivatives of this kind have been identified by a number of investigators from a variety of insertion mutations (10)(11)(12)(13)(14)(15)(16). In addition to changes in the frequency and timing of somatic reversion, these derivatives can involve alterations in the basal level of expression of the affected gene (9, 10, 16) and differences in the rate ofgerminal change (10,12,15).Two main hypotheses have been proposed to explain the molecular basis for changes in state of unstable mutations. The composition hypothesis (2, 17...