The luminal and discoid vacuole membranes of the superficial cell layer of the transitional epithelium of the mammalian urinary bladder have been studied by thin-sectioning and freeze-fracture-etch (FFE) electron microscope methods. For the FFE studies membranes were deposited on a cationized glass surface, covered by a thin copper disc, and fractured under liquid N2. Specimens were etched at -100°C and replicated at -190°C. A model of the lattice membrane derived from thin sections was used to predict the heights of the fracture faces above the glass surface . A hexagonal pattern of globular intramembrane particles spaced 160 A apart was seen in the external fracture (EF) face plaques as previously described and regarded as the dominant structure. However, very extensive areas of another pattern, seen before in only limited areas, have been found in the EF faces . The pattern consists of a smooth hexagonal lattice with the same space constant as the globular one but a different structure . By image analysis it consists of overlapping domains bordered by shared but incomplete metal rims . Each domain has a central spot of metal encircled by a shadow . The surface of the smooth lattice is partly complementary to the corresponding protoplasmic fracture (PF) face which shows a similar hexagonal lattice with the same space constant . The height of the smooth EF lattice above the glass substrate is the same as the plane of the center of the lipid bilayer predicted by the model . The mean heights of the particles of the globular EF lattice are greater than the total thickness of the membrane as predicted by the model and confirmed by measurements . The globular EF lattice is not complementary and it is concluded that the globular particles do not exist in the native membrane but arise artifactually during the preparatory procedures .Porter and Bonneville (36) first noted the unusual features of the luminal membrane of the mammalian urinary bladder epithelial cell, and it subsequently has been the subject of numerous investigations by groups associated with Porter (11,12,33,37,47) and 29,42,56,57), by ourselves (30,38,53,54), and others (28) . It is now established that 70-75% of the surface area of the luminal membrane and the membranes of the related discoid vacuoles in the luminal epithelial cell display external plaques 0.2-0.5 ,um in diameter comprised of particles in hexagonal array with p6 plane group symmetry spaced -160 A center-to-center . In negatively membranes are more flexible and are often sharply folded, particularly in the discoid vacuoles . For this reason the interplaque membrane is frequently called the "hinge" membrane to distinguish it from the "plaque" or "lattice" membrane .In freeze-fracture-etch (FFE) preparations the 120 A surface particles have a counterpart in an array of globular particles in the external fracture (EF) faces (8, 42). The EF particles usually appear globular and have been said by Staehelin et al . (47) to rise 40 A above the EF face and to fit into corresponding p...