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Overview of the model: PIP3 domains first expand, capturing membrane into the cup as the wave of actin polymerization moves outward; then slow and stall with the actin polymerization now remaining under the same area of membrane. The cup deepens and closes either at the lip or base depending on whether the zone of actin polymerization turns inward or not. Actin polymerization under the membrane causes an outward force (green arrows) and an opposite reaction force that can be transmitted to the membrane of the cup by retrograde flow of F-actin (blue arrows); the tension force creates a force whose direction depends on curvature and the elastic tension gradient (black arrows).

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