The function and mechanism of convergent extension during gastrulation of Xenopus laevis

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Abstract
The processes thought to function in Xenopus gastrulation include bottle cell formation, migration of cells on the roof of the blastocoel, and autonomous convergent extension of the circumblastoporal region. A review of recent and classical results shows that only the last accounts for the bulk of the tissue displacement of gastrulation, including spreading of the marginal zone toward the blastopore, involution of the marginal zone, and closure of the blastopore. Microsurgical manipulation and explantation studies, analysed by time-lapse video and cine microscopy, shows that the dorsal circumblastoporal region contains two regions which show either autonomous or semiautonomous convergent extension. The dorsal involuting marginal zone (IMZ) undergoes convergence (narrowing) and extension (lengthening) after its involution, beginning at the midgastrula stage and continuing through neurulation, such that it simultaneously extends posteriorly across the yolk plug and narrows the blastoporal circumference. Concurrently, the corresponding region of the overlying non-involuting marginal zone (NIMZ) begins a complementary convergent extension, but at a greater rate, which spreads vegetally to occupy surface area vacated by the IMZ. Tissue recombination experiments show that the deep cells of the dorsal IMZ bring about convergent extension. Labelling of small populations of these cells with a cell lineage tracer shows that convergent extension involves intercalation of deep cells to form a longer, narrower array. Direct time-lapse video and cine micrography of deep cells in cultured explants show that convergent extension involves radial and circumferential intercalation. Removal of the entire blastocoel roof of the early gastrula, including all or part of the NIMZ, shows that convergent extension of the IMZ alone can bring about its involution and blastopore closure. The role of convergent extension in gastrulation of other amphibians and other metazoans and its significance to related problems in early development are discussed.