Who's really in charge: Diverse follower cell behaviors in collective cell migration

Collective cell migrations drive morphogenesis, wound healing, and cancer dissemination. Cells located at the front are considered leaders while those behind them are defined topologically as followers. Leader cell behaviors, including chemotaxis and their coupling to followers, have been well-studied and reviewed. However, the contributions of follower cells to collective cell migration represent an emerging area of interest. In this perspective, we highlight recent research into the broadening array of follower cell behaviors found in moving collectives. We describe examples of follower cells that possess cryptic leadership potential and followers that lack that potential but contribute in diverse and sometimes surprising ways to collective movement, even steering from behind. We highlight collectives in which all cells both lead and follow, and a few passive passengers. The molecular mechanisms controlling follower cell function and behavior are just emerging and represent an exciting frontier in collective cell migration research.