Universal vortex formation time of flapping flight

Y Yukun Sun (Department of Biological and Environmental Engineering) E Emily Palmer (Department of Physics) C Christopher Dougherty (Department of Biological and Environmental Engineering) C Cade Sbrocco (Department of Physics) A Aspen Shih (Department of Biological and Environmental Engineering) J Jena Shields (Department of Applied and Engineering Physics) C Chris Roh (Department of Biological and Environmental Engineering)

Abstract

Biological flyers periodically flap their appendages to generate aerodynamic forces. Extensive studies have made significant progress in explaining the physics behind their propulsion in cruising by developing scaling laws of their flight kinematics. Notably Strouhal number ( St ; ratio of flapping frequency times stroke amplitude to cruising speed) has been found to fall in a narrow range for animal cruising flights. However, St exhibits strong correlation to flight conditions; as such, its universality has been confined to preferred flight conditions. Since the leading-edge vortices (LEV) on flapping appendages generate the majority of propulsive forces, here we take the perspective of LEV circulation maximization, which generalizes the dimensionless vortex formation time to flapping flight. The generalized vortex formation time scales the duration of vorticity injection with the rate of total vorticity growth inside the LEV and the maximum vorticity allowed inside it. By comparing the new scaling with St of previously reported animal cruising flights of 28 species, we show that the generalized vortex formation time is consistent across different animals and cruising locomotion, independent of flight conditions. This finding advances the fundamental principles underlying the complex wing kinematics of biological flyers and highlights a unifying framework for understanding biolocomotion.

Article Details

Volume / Issue Vol. 122, Issue 35
Published September 02, 2025
ISSN 0027-8424
Publisher National Academy of Sciences

Authors (7)

Y

Yukun Sun

Department of Biological and Environmental Engineering

E

Emily Palmer

Department of Physics

C

Christopher Dougherty

Department of Biological and Environmental Engineering

C

Cade Sbrocco

Department of Physics

A

Aspen Shih

Department of Biological and Environmental Engineering

J

Jena Shields

Department of Applied and Engineering Physics

C

Chris Roh

Department of Biological and Environmental Engineering