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1. Closer Appendage Spacing Augments Metachronal Swimming Speed by Promoting Tip Vortex Interactions

   https://doi.org/10.1093/icb/icab112  

   Ford, Mitchell P; Santhanakrishnan, Arvind (May 2021, Integrative and Comparative Biology)

   null (Ed.)

   Abstract Numerous species of aquatic invertebrates, including crustaceans, swim by oscillating multiple closely spaced appendages. The coordinated, out-of-phase motion of these appendages, known as “metachronal paddling,” has been well-established to improve swimming performance relative to synchronous paddling. Invertebrates employing this propulsion strategy cover a wide range of body sizes and shapes, but the ratio of appendage spacing (G) to the appendage length (L) has been reported to lie in a comparatively narrow range of 0.2 < G/L ≤ 0.65. The functional role of G/L on metachronal swimming performance is unknown. We hypothesized that for a given Reynolds number and stroke amplitude, hydrodynamic interactions promoted by metachronal stroke kinematics with small G/L can increase forward swimming speed. We used a dynamically scaled self-propelling robot to comparatively examine swimming performance and wake development of metachronal and synchronous paddling under varying G/L, phase lag, and stroke amplitude. G/L was varied from 0.4 to 1.5, with the expectation that when G/L is large, there should be no performance difference between metachronal and synchronous paddling due to a lack of interaction between vortices that form on the appendages. Metachronal stroking at nonzero phase lag with G/L in the biological range produced faster swimming speeds than synchronous stroking. As G/L increased and as stroke amplitude decreased, the influence of phase lag on the swimming speed of the robot was reduced. For smaller G/L, vortex interactions between adjacent appendages generated a horizontally oriented wake and increased momentum fluxes relative to larger G/L, which contributed to increasing swimming speed. We find that while metachronal motion augments swimming performance for closely spaced appendages (G/L <1), moderately spaced appendages (1.0 ≤ G/L ≤ 1.5) can benefit from the metachronal motion only when the stroke amplitude is large. 

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2. Hybrid Metachronal Rowing Augments Swimming Speed and Acceleration via Increased Stroke Amplitude

   https://doi.org/10.1093/icb/icab141  

   Ford, Mitchell P; Ray, William J; DiLuca, Erika M; Patek, S N; Santhanakrishnan, Arvind (June 2021, Integrative and Comparative Biology)

   null (Ed.)

   Synopsis Numerous aquatic invertebrates use drag-based metachronal rowing for swimming, in which closely spaced appendages are oscillated starting from the posterior, with each appendage phase-shifted in time relative to its neighbor. Continuously swimming species such as Antarctic krill generally use “pure metachronal rowing” consisting of a metachronal power stroke and a metachronal recovery stroke, while burst swimming species such as many copepods and mantis shrimp typically use “hybrid metachronal rowing” consisting of a metachronal power stroke followed by a synchronous or nearly synchronous recovery stroke. Burst swimming organisms need to rapidly accelerate in order to capture prey and/or escape predation, and it is unknown whether hybrid metachronal rowing can augment acceleration and swimming speed compared to pure metachronal rowing. Simulations of rigid paddles undergoing simple harmonic motion showed that collisions between adjacent paddles restrict the maximum stroke amplitude for pure metachronal rowing. Hybrid metachronal rowing similar to that observed in mantis shrimp (Neogonodactylus bredini) permits oscillation at larger stroke amplitude while avoiding these collisions. We comparatively examined swimming speed, acceleration, and wake structure of pure and hybrid metachronal rowing strategies by using a self-propelling robot. Both swimming speed and peak acceleration of the robot increased with increasing stroke amplitude. Hybrid metachronal rowing permitted operation at larger stroke amplitude without collision of adjacent paddles on the robot, augmenting swimming speed and peak acceleration. Hybrid metachronal rowing generated a dispersed wake unlike narrower, downward-angled jets generated by pure metachronal rowing. Our findings suggest that burst swimming animals with small appendage spacing, such as copepods and mantis shrimp, can use hybrid metachronal rowing to generate large accelerations via increasing stroke amplitude without concern of appendage collision. 

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3. Metachronal Swimming of Mantis Shrimp: Kinematics and Interpleopod Vortex Interactions

   https://doi.org/10.1093/icb/icab052  

   Garayev, Kuvvat; Murphy, David W (May 2021, Integrative and Comparative Biology)

   Abstract 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. 

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4. Exploring the swimming performance and the physical mechanisms of Tomopteris locomotion

   https://doi.org/10.1088/1748-3190/adad26  

   Battista, Nicholas A (February 2025, Bioinspiration & Biomimetics)

   Abstract Tomopterids are mesmerizing holopelagic swimmers. They use two modes of locomotion simultaneously: drag-based metachronal paddling and bodily undulation.Tomopterishas two rows of flexible, leg-like parapodia positioned on opposite sides of its body. Each row metachronally paddles out of phase to the other. Both paddling behaviors occur in concert with a lateral bodily undulation. However, when looked at independently, each mode appears in tension with the other. The direction of the undulatory wave is opposite of what one may expect for forward (FWD) swimming and appears to actively work act against the direction of swimming initiated by metachronal paddling. To investigate how these two modes of locomotion synergize to generate effective swimming, we created a self-propelled, fluid-structure interaction model of an idealizedTomopteris. We holistically explored swimming performance over a 3D mechanospace comprising parapodia length, paddling amplitude, and undulatory amplitude using a machine learning framework based on polynomial chaos expansions. Although undulatory amplitude minimally affected FWD swimming speeds, it helped mitigate the larger costs of transport that arise from either using more mechanically expensive (larger) paddling amplitudes and/or having longer parapodia. 

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5. The role of suction thrust in the metachronal paddles of swimming invertebrates

   https://doi.org/10.1038/s41598-020-74745-y  

   Colin, Sean P.; Costello, John H.; Sutherland, Kelly R.; Gemmell, Brad J.; Dabiri, John O.; Du Clos, Kevin T. (October 2020, Scientific Reports)

   Abstract An abundance of swimming animals have converged upon a common swimming strategy using multiple propulsors coordinated as metachronal waves. The shared kinematics suggest that even morphologically and systematically diverse animals use similar fluid dynamic relationships to generate swimming thrust. We quantified the kinematics and hydrodynamics of a diverse group of small swimming animals who use multiple propulsors, e.g. limbs or ctenes, which move with antiplectic metachronal waves to generate thrust. Here we show that even at these relatively small scales the bending movements of limbs and ctenes conform to the patterns observed for much larger swimming animals. We show that, like other swimming animals, the propulsors of these metachronal swimmers rely on generating negative pressure along their surfaces to generate forward thrust (i.e., suction thrust). Relying on negative pressure, as opposed to high pushing pressure, facilitates metachronal waves and enables these swimmers to exploit readily produced hydrodynamic structures. Understanding the role of negative pressure fields in metachronal swimmers may provide clues about the hydrodynamic traits shared by swimming and flying animals. 

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