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During angiogenesis, endothelial cells compete for the tip position duringangiogenesis: a phenomenon named tip cell overtaking. It is still unclear towhat extent tip cell overtaking is a side effect of sprouting or to what extenta biological function. To address this question, we studied tip cell overtakingin two existing cellular Potts models of angiogenic sprouting. In these modelsangiogenic sprouting-like behavior emerges from a small set of plausible cellbehaviors and the endothelial cells spontaneously migrate forwards andbackwards within sprouts, suggesting that tip cell overtaking might occur as aside effect of sprouting. In accordance with experimental observations, in oursimulations the cells' tendency to occupy the tip position can be regulatedwhen two cell lines with different levels of Vegfr2 expression are contributingto sprouting (mosaic sprouting assay), where cell behavior is regulated by asimple VEGF-Dll4-Notch signaling network. Our modeling results suggest that tipcell overtaking occurs spontaneously due to the stochastic motion of cellsduring sprouting. Thus, tip cell overtaking and sprouting dynamics may beinterdependent and should be studied and interpreted in combination.VEGF-Dll4-Notch can regulate the ability of cells to occupy the tip cellposition, but only when cells in the simulation strongly differ in their levelsof Vegfr2. We propose that VEGF-Dll4-Notch signaling might not regulate whichcell ends up at the tip, but assures that the cell that randomly ends up at thetip position acquires the tip cell phenotype.

Tip cell overtaking occurs as a side effect of sprouting in computational models of angiogenesis