Collective swimming pattern and synchronization of fish pairs (Gobiocypris rarus) in response to flow with different velocities

Collective behaviors in moving fish originate from social interactions, which are thought to be driven by beneficial factors, such as predator avoidance and reduced energy expenditure. Despite numerical simulations and physical experiments aiming at the hydrodynamic mechanisms and interaction rules, how shoaling is influenced by flow velocity and group size is still only partially understood. In this study, spatial distributions, kinematics, and synchronization states between pairs (smallest subsystem of a shoal) of Gobiocypris rarus were investigated in a recirculating swim tunnel with increasing flow velocities from 0.1 to 0.5 m/s (U_crit = 0.6 m/s). Tests of single fish were also conducted as the control group. The results of spatial distributions showed that fish pairs preferred to swim in the side-by-side configuration under high flows, while under low flows the neighboring fish's positions were more uniformly distributed around the focal fish in the transverse direction. Kinematic analysis revealed that fish pairs adopted similar tail beats (i.e., frequency and Strouhal number) as single fish in low flows, while in high flows both the frequency and Strouhal number of fish pairs were slightly lower. Moreover, the synchronization rates of fish pairs were found to increase with flow velocities, suggesting that synchronized swimming may be beneficial, especially in high flows.