The maize Suppressor-mutator (Spm) element can exist in one of three heritable forms: (1) a stably active form, (2) a stably inactive form, termed cryptic, and (3) a labile form, here termed programmable, in which the element exhibits one of a variety of heritable developmental programs of expression. Active elements are transcribed and are hypomethylated at sites upstream of the transcription start site, whereas inactive elements are transcriptionally silent and largely methylated at the upstream sites. Active (both stable and programmable), inactive programmable, and cryptic elements are unmethylated, partially methylated, and fully methylated, respectively, at sites within an 0.35-kb 80% G + C region just downstream from the transcription start site. An active Spin element in a genome with a cryptic element promotes its partial demethylation but not its transcriptional activation. In contrast, a trans-acting Spin promotes extensive demethylation and transcriptional activation of an inactive programmable element, as well as its heritable reactivation. These observations define the molecular components of the Spin element's developmental regulatory mechanism. We discuss their general relevance to the developmental regulation of gene expression.[ 1957, 1958, 1959, 1965aPeterson 1966;Fowler and Peterson 1978). These include the cryptic form, in which the element is genetically silent. Both McClintock and Peterson also identified elements that were capable of undergoing reversible transitions between the active and inactive phases in a characteristic and heritable pattern during development (McClintock 1957(McClintock , 1958(McClintock , 1962(McClintock , 1965a(McClintock , b, 1971Peterson 1966;Fowler and Peterson 1978). We showed recently that the genetic mechanism responsible for the developmental control of Spin expression has two components, which we designated the phase setting and the phase program (Fedoroff and Banks 1988). The phase setting determines whether the element is genetically active or inactive. The phase program determines the heritability of the phase setting, as well as when, where, and how frequently it will be reversed during plant development. The underlying genetic mechanism is heritable, reversible, and capable of promoting the incremental transition of the element from a stably active to a stably inactive (cryptic) form, implying the involvement of multiple epigenetic events (Fedoroff and Banks 1988). In view of the accumulating evidence that the inactivation of gene expression, in general (for reviews, see Razin et al. 1984;Adams and Burdon 1985;Holliday 1987;Cedar 1988), and maize transposable elements, in particular Bennetzen 1985;Chandler and Walbot 1986;Schwartz and Dennis 1986;Chomet et al. 1987; Fedoroff et al. 1988a, b)