The organization of septate junctions during morphogenesis of imaginal disks is described from freeze-fracture replicas and thin sections with a view to understanding junction modulation during rearrangements of cells in epithelia. The septate junctions of each epithelial cell of the disk are distributed in a number of discrete domains equal to the number of neighboring cells. Individual septa traverse domains of contact between pairs of adjacent cells, turn downwards at the lateral boundary of the domain and run parallel to the intersection with a third cell. This arrangement leaves small channels at three-cell intersections that are occupied by specialized structures termed "tricellular plugs." Cell rearrangement involves a progressive change in the width of contact domains between adjacent cells, until old contacts are broken and new ones established. It is proposed that the septate junction adjusts to the changing width of domains by the compaction or extension of existing septa. This redistribution of septa theoretically allows a transepithelial barrier to be maintained during cell rearrangements. The applicability of this model to other epithelial tissues is discussed.The morphogenesis of an epithelium has previously been assumed to be limited to movements of entire epithelial sheets, with each cell within the epithelium retaining its immediate neighbors (1, 2). The presence of specialized connections between adjacent epithelial cells and the absence of morphological evidence of locomotion undoubtedly contributed to this view. However, rearrangements of cells (i.e., cell movements involving change of neighbors within a contiguous epithelial sheet) have recently been described during the evagination of imagined disks in Drosophila (3,4), neurulation in newts (5), gastrulation in Xenopus (6), regeneration in Hydra (7), and epiboly in teleosts (8). Thus, cell rearrangement appears to be a widespread phenomenon in epithelial morphogenesis.In most of the examples cited above, intercellular junctions persist during cell rearrangement (3,(6)(7)(8). Because junctions are sites of intercellular adhesion between pairs of adjacent cells, the exchange of neighbors requires some form ofjnnction modulation (9). Tight junctions of vertebrates and septate junctions of invertebrates present particularly formidable bartiers to cell rearrangement because they bridge or close off the intercellular space between a given cell and all of its neighbors, ensuring contiguity of the epithelial layer and providing a barrier to the extracellular movement of molecules across the epithelium (10-12). Thus, each cell appears to be fixed with respect to its neighbors. This paper focuses on the distribution of septa and the potentially dynamic organization of the septate junction during cell rearrangement in imagined disks. The detailed structure of the septate junction is well known. Rows of septa connect adjacent ceils and separate the apical from the basal intercellular space. The plasma membranes on both sides of a septum conta...