Flow structure and transport characteristics of feeding and exchange currents generated by upside-down Cassiopea jellyfish.

Quantifying the flows generated by the pulsations of jellyfish bells is crucial for understanding the mechanics and efficiency of their swimming and feeding. Recent experimental and theoretical work has focused on the dynamics of vortices in the wakes of swimming jellyfish with relatively simple oral arms and tentacles. The significance of bell pulsations for generating feeding currents through elaborate oral arms and the consequences for particle capture are not as well understood. To isolate the generation of feeding currents from swimming, the pulsing kinematics and fluid flow around the benthic jellyfish Cassiopea spp. were investigated using a combination of videography, digital particle image velocimetry and direct numerical simulation. During the rapid contraction phase of the bell, fluid is pulled into a starting vortex ring that translates through the oral arms with peak velocities that can be of the order of 10 cm s(-1). Strong shear flows are also generated across the top of the oral arms throughout the entire pulse cycle. A coherent train of vortex rings is not observed, unlike in the case of swimming oblate medusae such as Aurelia aurita. The phase-averaged flow generated by bell pulsations is similar to a vertical jet, with induced flow velocities averaged over the cycle of the order of 1-10 mm s(-1). This introduces a strong near-horizontal entrainment of the fluid along the substrate and towards the oral arms. Continual flow along the substrate towards the jellyfish is reproduced by numerical simulations that model the oral arms as a porous Brinkman layer of finite thickness. This two-dimensional numerical model does not, however, capture the far-field flow above the medusa, suggesting that either the three-dimensionality or the complex structure of the oral arms helps to direct flow towards the central axis and up and away from the animal.