Synopsis: Metachronal motion is used across a wide range of organisms for a diverse set of functions. However, despite its ubiquity, analysis of this behavior has been difficult to generalize across systems. Here we provide an overview of known commonalities and differences between systems that use metachrony to generate fluid flow. We also discuss strategies for standardizing terminology and defining future investigative directions that are analogous to other established subfields of biomechanics. Finally, we outline key challenges that are common to many metachronal systems, opportunities that have arisen due to the advent of new technology (both experimental and computational), and next steps for community development and collaboration across the nascent network of metachronal researchers.

The present work is devoted to the analysis of drop impact on a deep liquid pool, focusing on the high-energy splashing regimes caused by large raindrops at high velocities. Such cases are characterized by short time scales and complex mechanisms, thus they have received very little attention until now. The BASILISK open-source solver is used to perform three-dimensional direct numerical simulations. The capabilities of octree adaptive mesh refinement techniques enable capturing of the small-scale features of the flow, while the volume of fluid approach combined with a balanced-force surface-tension calculation is applied to advect the volume fraction of the liquids and reconstruct the interfaces. The numerical results compare well with experimental visualizations: both the evolution of crown and cavity, the emanation of ligaments, the formation of bubble canopy and the growth of a downward-moving spiral jet that pierces through the cavity bottom, are correctly reproduced. Reliable quantitative agreements are also obtained regarding the time evolution of rim positions, cavity dimensions and droplet distributions through an observation window. Furthermore, simulation gives access to various aspects of the internal flows, which allows us to better explain the observed physical phenomena. Details of the early-time dynamics of bubble ring entrapment and splashing performance, the formation/collapse of bubble canopy and the spreading of drop liquid are discussed. The statistics of droplet size show the bimodal distribution in time, corroborating distinct primary mechanisms of droplet production at different stages.

Mantis shrimp swim via metachronal rowing, a pattern in which the pleopods (swimming limbs) stroke sequentially, starting with the last pair and followed by anterior neighbors. A similar swimming pattern is used at various sizes, Reynolds numbers, and advance ratios by diverse organisms including ciliates, ctenophores, copepods, krill, and lobsters. Understanding this type of locomotion is important because it is widespread and may inspire the design of underwater vehicles where efficiency, robustness, and maneuverability are desired. However, detailed measurements of the flow around free-swimming, metachronally rowing organisms are scarce, especially for organisms swimming in a high Reynolds number regime (Re ≥ 10⁴). In this study, we present time-resolved, planar PIV measurements of a swimming peacock mantis shrimp (Odontodactylus scyllarus). Simultaneous kinematics measurements of the animal, which had body and pleopod lengths of 114 and 20 mm, respectively, reveal mean swimming speeds of 0.2–1.9 m s⁻¹ and pleopod beat frequencies of 3.6–13 Hz, corresponding to advance ratios of 0.75–1.84 and body-based Reynolds numbers of 23,000–217,000. Further, the animal’s stroke is not purely metachronal, with a long phase lag between initiation of the first and fifth pleopod power strokes. Flow measurements in the sagittal plane show that each stroking pleopod pair creates a posteriorly moving tip vortex which evades destruction by the recovery strokes of other pleopod pairs. The vortex created by the anteriormost pleopod pair is the strongest and, owing to the animal’s high advance ratio, is intercepted by the power stroke of the posteriormost pleopod pair. The vortex strength increases as a result of this interaction, which may increase swimming speed or efficiency. A relationship for vortex interception by the posterior pleopod is proposed that relates the phase lag between the interacting pleopods to the beat frequency, distance between those pleopods, and speed of the vortex relative to the animal. We describe this interaction with a novel parameter called the interpleopod vortex phase matching Strouhal number $StIVPM$ which is equal to the phase lag between interacting pleopods. This new nondimensional parameter may be useful in predicting the conditions where a constructive interaction may occur in other species or in physical models. Finally, we relate the advance ratio to the Reynolds number ratio, the ratio between the body-based Reynolds number and the pleopod-based Reynolds number. The importance of these parameters in promoting the interpleopod vortex interactions identified here, in dynamically scaled experiments, and in wake signatures behind schooling metachronal swimmers is discussed.

In drag-based swimming, individual propulsors operating at low Reynolds numbers (where viscous forces dominate over inertial forces) must execute a spatially asymmetric stroke to produce net fluid displacement. Temporal asymmetry (that is, differing duration between the power vs. recovery stroke) does not affect the overall generated thrust in this time-reversible regime. Metachronal rowing, in which multiple appendages beat sequentially, is used by a wide variety of organisms from low to intermediate Reynolds numbers. At the upper end of this range, inertia becomes important, and increasing temporal asymmetry can be an effective way to increase thrust. However, the combined effects of spatial and temporal asymmetry are not fully understood in the context of metachronal rowing. To explore the role of spatiotemporal asymmetry in metachronal rowing, we combine laboratory experiments and reduced-order analytical modeling. We measure beat kinematics and generated flows in two species of lobate ctenophores across a range of body sizes, from 7 to 40 mm in length. We observe characteristically different flows in ctenophores of differing body size and Reynolds number, and a general decrease in spatial asymmetry and increase in temporal asymmetry with increasing Reynolds number. We also construct a one-dimensional mathematical model consisting of a row of oscillating flat plates whose flow-normal areas change with time, and use it to explore the propulsive forces generated across a range of Reynolds numbers and kinematic parameters. The model results show that while both types of asymmetry increase force production, they have different effects in different regions of the parameter space. These results may have strong biological implications, as temporal asymmetry can be actively controlled while spatial asymmetry is likely to be partially or entirely driven by passive fluid-structure interaction.