Surface Specializations of Fundulus Cells and Their Relation to Cell Movements during Gastrulation

Cell movements in Fundulus blastoderms during gastrulation were studied utilizing time-lapse cinemicrography and electron microscopy. Time-lapse films reveal that cells of the enveloping layer undulate and sometimes separate briefly but remain together in a cohesive layer. During epiboly, the marginal enveloping layer cells move over the periblast as it expands over the yolk sphere. Movement occurs as a result of ruffled membrane activity of the free borders of the marginal cells. Deep blastomeres become increasingly active during blastula and gastrula stages. Lobopodia project from the blastomeres in blastulae and adhere to other cells in gastrulae, giving the cells traction for movement. Contact specializations are formed by the lateral adjacent plasma membranes of enveloping layer cells. An apical junction is characterized by an intercellular gap of 60-75 A. Below this contact, the plasma membranes are separated by 120 A or more. In mid-gastrulae, cytoplasmic fibrils occur adjacent to some apical junctions, and small desmosomes appear below the apical junction. Septate desmosomes also appear at this time. A junction with an intercellular gap of 60 A occurs between marginal enveloping layer cells and periblast. Contacts between deep blastomeres become numerous in gastrulae and consist of contacts at the crests of surface undulations, short areas of contact in which the plasma membranes are 60 or 120 A apart, and long regions characterized by a 200-A intercellular gap. Lobopodia contact other blastomeres only in gastrulae. These junctions contain a 200-A intercellular space. Some deep blastomeres are in contact with the tips of periblast microvilli. The mechanism of epiboly in Fundulus is discussed and reevaluated in terms of these observations. The enveloping layer is adherent to the margin of the periblast and moves over it as a coherent cellular sheet. Periblast epiboly involves a controlled flow of cytoplasm from the thicker periblast into the thinner yolk cytoplasmic layer with which it is continuous. Deep cells move by adhering to each other, to the inner surface of the enveloping layer, and to the periblast.