Hummingbirds rapidly respond to the removal of vision and control a sequence of rate-commanded maneuvers in milliseconds

Hummingbird flight is the epitome of extreme aerial agility and controlled stability, as hummingbirds routinely exercise a variety of stunning aerobatic feats. Yet, the control of these amazing maneuvers is not well understood. Here we examined how hummingbirds control a sequence of maneuvers within milliseconds and tested whether and when their vision is active during this rapid process. We elicited escape flight in calliope hummingbirds and removed visible light at various instants during the maneuvers and quantified their flight kinematics and responses. We show that the escape maneuvers were composed of rapidly-controlled sequential modules, including evasion, reorientation, nose-down dive, forward flight, and nose-up to hover. The hummingbirds did not respond to the light removal during evasion and reorientation until a critical light-removal time; afterward they showed two categories of luminance-based responses that rapidly altered maneuvering modules to terminate the escape. We also show that hummingbird maneuvers are rate-commanded and require no active braking, i.e., their body angular velocities were proportional to the change of wing motion patterns, a trait that likely alleviates the computational demand on flight control. Together, this work uncovers the key traits of hummingbird agility, which can also inform and inspire designs for next-generation agile aerial systems.