The mechanism of chick blastoderm expansion.

At the time of laying, the domestic fowl blastoderm measures 4 mm across. After 4 days' incubation, the extra-embryonic yolk-sac tissues have expanded to encompass the whole yolk mass. This expansion involves the migration over the inner surface of the vitelline membrane of a specialized band of 'edge cells' at the blastoderm periphery. As they move, they pull out the blastoderm behind them, setting up a considerable tension. Expansion also involves cell proliferation and changes in cell shape. This paper attempts to show how locomotion, tension, proliferation and changes in cell shape all contribute to the orderly process of expansion. As a simplification, only the extra-embryonic epiblast is considered here. The findings are: 1. Expansion does not occur at a constant rate, but starts slowly, rises to a peak (over 500 mum/h) at around 3 days, and then slows as coverage of the yolk mass nears completion. 2. During the first day of incubation, edge-cell migration produces a tension in the blastoderm. This rises to peak at 20-24 h, then declines. This tension may be due to an imbalance between expansion by migration and expansion by proliferation. 3. Migration of edge cells can be affected by tension in the blastoderm, i.e. very high tension may hold them back. However, the tension level normally found in the blastoderm seems not to do so. The low rate of expansion in the first day is therefore not due to the high level of tension. It may instead be due to changes in edge-cell organization. 4. Proliferation occurs throughout the extra-embryonic epiblast during the expansion period. It is not restricted to the blastoderm periphery. After the yolk has been covered, the epiblast continues to grow, with proliferation restricted largely to band just distal to the advancing edge of the area vasculosa. 5. Cell shape and arrangement change considerably during expansion. The epiblast of the unincubated embryo is a monolayer of tall cells. During expansion, these become considerably flattened so that each contributes a larger amount to yolk-sac surface area.