Swimming Strategies Research Articles

Encoding spatiotemporal asymmetry in artificial cilia with a ctenophore-inspired soft-robotic platform

A remarkable variety of organisms use metachronal coordination (i.e., numerous neighboring appendages beating sequentially with a fixed phase lag) to swim or pump fluid. This coordination strategy is used by microorganisms to break symmetry at small scales where viscous effects dominate and flow is time-reversible. Some larger organisms use this swimming strategy at intermediate scales, where viscosity and inertia both play important roles. However, the role of individual propulsor kinematics-especially across hydrodynamic scales-is not well-understood, though the details of propulsor motion can be crucial for the efficient generation of flow. To investigate this behavior, we developed a new soft robotic platform using magnetoactive silicone elastomers to mimic the metachronally coordinated propulsors found in swimming organisms. Furthermore, we present a method to passively encode spatially asymmetric beating patterns in our artificial propulsors. We investigated the kinematics and hydrodynamics of three propulsor types, with varying degrees of asymmetry, using Particle Image Velocimetry and high-speed videography. We find that asymmetric beating patterns can move considerably more fluid relative to symmetric beating at the same frequency and phase lag, and that asymmetry can be passively encoded into propulsors via the interplay between elastic and magnetic torques. Our results demonstrate that nuanced differences in propulsor kinematics can substantially impact fluid pumping performance. Our soft robotic platform also provides an avenue to explore metachronal coordination at the meso-scale, which in turn can inform the design of future bioinspired pumping devices and swimming robots.

Adult caudal fin shape is imprinted in the embryonic fin fold.

Appendage shape is formed during development (and re-formed during regeneration) according to spatial and temporal cues that orchestrate local cellular morphogenesis. The caudal fin is the primary appendage used for propulsion in most fish species, and exhibits a range of distinct morphologies adapted for different swimming strategies, however the molecular mechanisms responsible for generating these diverse shapes remain mostly unknown. In zebrafish, caudal fins display a forked shape, with longer supportive bony rays at the periphery and shortest rays at the center. Here, we show that a premature, transient pulse of sonic hedgehog a (shha) overexpression during late embryonic development results in excess proliferation and growth of the central rays, causing the adult caudal fin to grow into a triangular, truncate shape. Both global and regional ectopic shha overexpression are sufficient to alter fin shape, and forked shape may be rescued by subsequent treatment with an antagonist of the canonical Shh pathway. The induced truncate fins show a decreased fin ray number and fail to form the hypural diastema that normally separates the dorsal and ventral fin lobes. While forked fins regenerate their original forked morphology, truncate fins regenerate truncate, suggesting that positional memory of the fin rays can be permanently altered by a transient treatment during embryogenesis. Ray finned fish have evolved a wide spectrum of caudal fin morphologies, ranging from truncate to forked, and the current work offers insights into the developmental mechanisms that may underlie this shape diversity.

Use of Ordered Beta Regression Unveils Cognitive Flexibility Index and Longitudinal Cognitive Training Signatures in Normal and Alzheimer's Disease Pathological Aging.

Generalized linear mixed models (GLMMs) are a cornerstone data analysis strategy in behavioral research because of their robustness in handling non-normally distributed variables. Recently, their integration with ordered beta regression (OBR), a novel statistical tool for managing percentage data, has opened new avenues for analyzing continuous response data. Here, we applied this combined approach to investigate nuanced differences between the 3xTg-AD model of Alzheimer's disease (AD) and their C57BL/6 non-transgenic (NTg) counterparts with normal aging in a 5-day Morris Water Maze (MWM) test protocol. Our longitudinal study included 22 3xTg-AD mice and 15 NTg mice (both male and female) assessed at 12 and 16 months of age. By identifying and analyzing multiple swimming strategies during three different paradigms (cue, place task, and removal), we uncovered genotypic differences in all paradigms. Thus, the NTg group exhibited a higher percentage of direct search behaviors, while an association between circling episodes and 3xTg-AD animals was found. Furthermore, we also propose a novel metric-the "Cognitive Flexibility Index"-which proved sensitive in detecting sex-related differences. Overall, our integrated GLMMs-OBR approach provides a comprehensive insight into mouse behavior in the MWM test, shedding light on the effects of aging and AD pathology.

Swimming behaviour of downstream migrating carp in accelerating flows

The downstream migration of fish is impacted by accelerating flow. This study coupled computational fluid dynamics (CFD) modelling and laboratory observations to investigate the effects of two different accelerating flow hydrodynamics on the swimming behaviours of two fish species. The silver carp (Hypophthalmichthys molitrix) and bighead carp (Hypophthalmichthys nobilis) were chosen as target fish species and four hydrodynamic parameters were selected as main indicators to affect fish swimming behaviours. Four specific swimming behaviours were classified in the contraction flume based on fish downstream trajectories. The results indicated that high accelerating flow, low turbulent kinetic energy (Tke ) and shear stress generated the symmetric distribution of fish along two boundaries of the contraction flume. It also indicated that discharge increase would increase the fish entrainment risk from intakes to diversion pipe, while two fish species exhibited different swimming behaviour patterns. Velocity and velocity gradient had more significant effects than other hydrodynamic parameters on fish swimming strategies. There was mutual compensation among fish tail-beat frequency, angle and amplitude. The results illustrated the relationship between the swimming behaviours and hydrodynamics, which could contribute to understanding the fish downstream migration behaviours in accelerating flow.

Nature-inspired miniaturized magnetic soft robotic swimmers

State-of-the-art biomedical applications such as targeted drug delivery and laparoscopic surgery are extremely challenging because of the small length scales, the requirements of wireless manipulation, operational accuracy, and precise localization. In this regard, miniaturized magnetic soft robotic swimmers (MSRS) are attractive candidates since they offer a contactless mode of operation for precise path maneuvering. Inspired by nature, researchers have designed these small-scale intelligent machines to demonstrate enhanced swimming performance through viscous fluidic media using different modes of propulsion. In this review paper, we identify and classify nature-inspired basic swimming modes that have been optimized over large evolutionary timescales. For example, ciliary swimmers like Paramecium and Coleps are covered with tiny hairlike filaments (cilia) that beat rhythmically using coordinated wave movements for propulsion and to gather food. Undulatory swimmers such as spermatozoa and midge larvae use traveling body waves to push the surrounding fluid for effective propulsion through highly viscous environments. Helical swimmers like bacteria rotate their slender whiskers (flagella) for locomotion through stagnant viscid fluids. Essentially, all the three modes of swimming employ nonreciprocal motion to achieve spatial asymmetry. We provide a mechanistic understanding of magnetic-field-induced spatiotemporal symmetry-breaking principles adopted by MSRS for the effective propulsion at such small length scales. Furthermore, theoretical and computational tools that can precisely predict the magnetically driven large deformation fluid–structure interaction of these MSRS are discussed. Here, we present a holistic descriptive review of the recent developments in these smart material systems covering the wide spectrum of their fabrication techniques, nature-inspired design, biomedical applications, swimming strategies, magnetic actuation, and modeling approaches. Finally, we present the future prospects of these promising material systems. Specifically, synchronous tracking and noninvasive imaging of these external agents during in vivo clinical applications still remains a daunting task. Furthermore, their experimental demonstrations have mostly been limited to in vitro and ex vivo phantom models where the dynamics of the testing conditions are quite different compared the in vivo conditions. Additionally, multi-shape morphing and multi-stimuli-responsive modalities of these active structures demand further advancements in 4D printing avenues. Their multi-state configuration as an active solid-fluid continuum would require the development of multi-scale models. Eventually, adding multiple levels of intelligence would enhance their adaptivity, functionalities, and reliability during critical biomedical applications.

Additive manufacturing inert gas flow path strategies for multi-laser powder bed fusion systems and their impact on lattice structure mechanical responses

Multi-laser Additive Manufacturing systems hold great potential to increase productivity. However, adding multiple energy sources to a powder bed fusion system requires careful selection of a laser scan and inert gas flow strategy to optimize component performance. In this work, we explore four different laser scan and argon flow strategies on the quasi-static compressive mechanical response of Body Centered Cubic lattices. Three strategies employ a swim lane method where laser pathing tends to progress parallel to argon flow. Method one only uses a single laser while method two uses four, both with the laser path working against the argon flow. The third method uses four lasers, each operating in their own lane like the second method, but the laser pathing progresses with the argon flow. The fourth method has all four lasers operating in quadrants and the laser pathing trends against the argon flow.The single-laser strategy generally had the lowest mechanical responses compared to the other three strategies. A quadrant strategy generally had the highest quasi-static mechanical responses and was at least 25% greater in stiffness, yield force, ultimate force, and energy absorption when compared to the single laser strategy. However, the four-laser swim strategy where the laser pathing tends against the argon flow was found to be statistically similar to the quadrant strategy. It is hypothesized that spatter introduced onto the powder layer from the melt pool and particle entrainment may be worse for laser pathing which trends with the argon flow direction. Additionally, the additional energy added to the build volume helps to mitigate inter-layer cool time which reduces temperature gradients. This shows that multi-laser AM systems have an impact on part performance and potentially shows lattices built with multi-laser AM systems may have certain advantages over single-laser AM systems.

Undulatory Propulsion at Milliscale on Water Surface.

The oscillatory pitch motion at the leading edge of a millimeter-scale flexible sheet on the water surface can generate undulatory locomotion for swimming, similar to a honeybee vibrating its wings for propulsion. The influence of various parameters on such swimming strategy remains unexplored. This study uses magnetic milliswimmers to probe the propulsion mechanics and impact of different parameters. It is found that this undulatory propulsion is driven by capillary forces and added mass effects related to undulatory waves of the milliswimmers, along with radiation stress stemming from capillary waves at the interface. Modifying the parameters such as actuation frequency, pitch amplitude, bending stiffness, and hydrofoil length alters the body waveform, thus, affecting the propulsion speed and energy efficiency. Although undulatory motion is not a prerequisite for water surface propulsion, optimizing body stiffness to achieve a proper undulatory waveform is crucial for efficient swimming, balancing energy consumption, and speed. The study also reveals that the induced water flow is confined near the water surface, and the flow structures evolve with varying factors. These discoveries advance the understanding of undulatory water surface propulsion and have implications for the optimal design of small-scale swimming soft robots in the future.

Morphology and motor behavior of endemic fishes in the upper reaches of the Yangtze River basin.

Dam construction alters the hydrodynamic conditions, consequently impacting the swimming behavior of fish. To explore the effect of flow hydrodynamics on fish swimming behavior, five endemic fish species in the upper Yangtze River basin were selected. Through high-speed video visualization and computer analysis, these species' swimming patterns under different flow velocities (0.1-1.2 m/s) were investigated. The kinematic and morphological characteristics of the fish were presented. The principal component analysis was used to analyse the main factors influencing the swimming ability of fish and to determine the correlation coefficients among fish behavior indicators. Fish exhibited three different swimming patterns under different flow velocities. Low velocity (0.1-0.3 m/s) corresponds to free motion, middle velocity (0.4-0.7 m/s) corresponds to cruising motion, and high velocity corresponds to stress motion (0.8-1.2 m/s). The fish kinematic index curves were obtained, and four of five fish species showed two extreme points, which means the optimal and adverse swimming strategies can be determined. With the increase in flow velocity, the tail-beat frequency showed an increasing trend, whereas the tail-beat angle and amplitude showed a decreasing trend. Morphological and kinematic parameters were the two main indexes that affect the swimming ability of fish, which accounts for 41.9% and 26.9%, respectively.

Effect of Total Dissolved Gas Supersaturation on Swimming Performance of Migratory Fish for Traversing Velocity Barriers

This study investigates the impact of total dissolved gas supersaturation (TDGS) on the swimming capabilities of migratory fish (S. prenanti), a common issue during high dam discharges in flood seasons. We assessed fish exposed to 130% TDGS for 2 hr, focusing on their swimming performance in a controlled environment. In our experiments, control group fish, utilizing prolonged swimming, showed reduced maximum distances as flow velocities increased from 3 to 10 BL/s (body length per second), covering distances between 1,285 and 119 BL. In contrast, TDGS‐exposed fish achieved only 15%–95% of these distances. Under burst swimming conditions, control group fish also demonstrated a decrease in maximum distances with increasing flow velocity, achieving 280–124 BL, while TDGS‐exposed fish reached just 48%–64% of these distances. Notably, the critical flow velocity (Ufcrit) for transitioning from prolonged to burst activity level was lower for the TDGS group (7.2 BL/s) compared with the control (9.8 BL/s). In open flume trials, TDGS‐exposed fish displayed a stronger upstream swimming inclination beyond Ufcrit, indicated by quicker times, higher speeds, and shorter trajectories. This study provides novel insights into the adaptive swimming strategies and flow velocity responses of fish under TDGS stress.

DISEÑOS DE MICRODISPOSITIVOS EFICIENTES PARA SEPARACIÓN DE COANOFLAGELADOS

Choanoflagellates are microorganisms of great interest in evolutionary biology because they are considered to be the closest living relatives to animals. In their unicellular form, they developed phenotypes with different swimming strategies and the ability to produce sexual gametes. Based on the work of Sparacino et al.,J. Phys. D: Appl. Phys.(2020), new designs of microfluidic devices were proposed and explored with consideration to both microfabrication restrictions and microscopy observation limitations. Through a phenomenological model, adapted to the dynamics of unicellular choanoflagellates in an asymmetric microdevice, it was possible to separate the fast cells, reconcentrating them up to 8times. It was observed in a temporal window independent of the geometry, minimizing the biological variation of the samples. Furthermore, new biotechnological applications were explored to model different biological behaviors related to the taxis responses experienced by a cell. It was noted that populations with lower change of direction are efficiently concentrated in the proposed microdevice

Exploring tadpole swimming strategies and dynamics

A model tadpole provides insights into efficient swimming patterns that could inspire future robotic microswimmers.

Search strategy analysis reveals sex‐specific deficits in spatial reference memory in 5xFAD mice

AbstractBackgroundSpatial disorientation is one of the earliest symptoms in Alzheimer’s disease. Search strategy classification systems to study navigational deficits in Alzheimer disease have been developed but aren’t widely available. Hence, we studied the navigational strategies in the 5xFAD murine model and evaluated differences between sexes.MethodWe studied spatial reference memory in 3‐month‐, 7‐month‐ and 12‐month‐old female and male 5xFAD mice and their aged‐matched controls in the Morris water maze. Furthermore, a detailed analysis of the swimming strategies using “Pathfinder” was performed. Navigational strategies were classified in spatial (“direct path”, “directed search”, “focal search”, “indirect search”) and non‐spatial categories (“chaining”, “scanning”, “random search”, thigmotaxis”).ResultEscape latency was prolonged in 5xFAD mice compared to WT, only at 7 and 12 months, with greater magnitude in females than males. Regarding search strategies during acquisition training, the most common spatial strategy was “indirect search” whereas “random search” was the most frequent non‐spatial strategy. At 3 months, no clear difference was seen between 5xFAD and WT animals. However, at 7 months, 5xFAD mice used more frequently non‐spatial strategies. Although transgenic mice progressively shifted to more spatial strategies as they learned (mainly “indirect search”), WT animals applied more frequently spatial strategies with higher cognitive value such as “direct path”, “directed search” and “focal search”. In comparison, 12 month‐old 5xFAD mice used more non‐spatial strategies, mainly “random search”, than the WT and their transition to “indirect search” occurred more rarely than in younger mice. Overall, all animals increased the use of non‐spatial strategies as they aged but this increment was greater and earlier in 5xFAD females than in 5xFAD males. Lastly, in the probe trial, 5xFAD female mice at 3 and 12 months applied “random search” more often, while the WT chose “indirect search” with higher frequency. No difference was seen in males during probe trial.ConclusionConventional behavior analysis of escape latencies revealed spatial reference memory deficits in 7m and 12m 5XFAD mice. Deficits in spatial memory were more pronounced in female 5XFAD mice than in male mice. 5XFAD mice, regardless of sex, used a qualitatively and quantitatively different search strategy pattern than WT animals.

First insights into the shortfin mako shark (Isurus oxyrinchus) fine-scale swimming behaviour.

As regional endotherms, lamnid sharks can sustain high cruising speeds and perform frequent speed bursts. However, since endothermy comes with high energetic costs, lamnids may adopt different swimming strategies to manage their energy budget. Understanding such strategies is essential to provide behavioural and physiological context to their broader movement ecology. The endangered shortfin mako (Isurus oxyrinchus) possibly has the highest energy requirements among lamnids, but our understanding of its swimming behaviour is still limited. We equipped three shortfin mako sharks with high-resolution multi-sensor tags to measure their swimming kinematics in the wild. While swimming horizontally, individuals favoured tail-beat frequencies around 0.6 Hz at speeds comparable to those of ectothermic sharks (ca 0.5 m s-1). All individuals displayed yo-yo-like diving patterns where, for a given tail-beat frequency, speeds were higher during descents, as expected for a negatively buoyant fish. Contrary to what was expected, gliding was almost absent (less than 1.31%). Speed bursts reaching up to 3.6 m s-1 were observed during the day but ceased shortly after dusk, implying a diel change in swimming behaviour. As large-scale research efforts are hindered by this species' increasing rarity, opportunistic high-resolution datasets, like the present, are fundamental to improve our understanding of shortfin mako's behaviour and ecology.

Fish swimming mode and body morphology affect the energetics of swimming in a wave-surge water flow.

Fish swimming modes and the shape of both the fins and body are expected to affect their swimming ability under different flow conditions. These swimming strategies and body morphologies often correspond to distributional patterns of distinct functional groups exposed to natural and variable water flows. In this study, we used a swimming-respirometer to measure energetic costs during prolonged, steady swimming and while station holding in a range of simulated oscillatory wave-surge water flows, within the natural range of flow speeds and wave frequencies on coral reefs. We quantified the net cost of swimming (NCOS, metabolic costs above resting) for four reef fish species with differences in swimming mode and morphologies of the fin and body: a body and caudal fin (BCF) swimmer, the Hawaiian flagtail, Kuhlia xenura, and three pectoral fin swimmers, the kole tang, Ctenochaetus strigosus, the saddle wrasse, Thalassoma duperrey, and the Indo-Pacific sergeant major, Abudefduf vaigiensis. We found that the BCF swimmer had the highest rates of increase in NCOS with increasing wave frequency (i.e. increased turning frequency) compared with the pectoral fin swimmers. The wrasse, with a more streamlined, higher body fineness, had lower rates of increase in NCOS with increasing swimming speeds than the low body fineness species, but overall had the highest swimming NCOS, which may be a result of a higher aerobic swimming capacity. The deep-bodied (low fineness) pectoral fin swimmers (A. vaigiensis and C. strigosus) were the most efficient at station holding in oscillating, wave-surge water flows.

Search Strategy Analysis of 5xFAD Alzheimer Mice in the Morris Water Maze Reveals Sex- and Age-Specific Spatial Navigation Deficits

Spatial disorientation and navigational impairments are not only some of the first memory deficits in Alzheimer's disease, but are also very disease-specific. In rodents, the Morris Water Maze is used to investigate spatial navigation and memory. Here, we examined the spatial memory in the commonly used 5xFAD Alzheimer mouse model in a sex- and age-dependent manner. Our findings show first spatial learning deficits in 7-month-old female 5xFAD and 12-month-old male 5xFAD mice, respectively. While the assessment of spatial working memory using escape latencies provides a global picture of memory performance, it does not explain how an animal solves a spatial task. Therefore, a detailed analysis of swimming strategies was performed to better understand the behavioral differences between 5xFAD and WT mice. 5xFAD mice used a qualitatively and quantitatively different search strategy pattern than wildtype animals that used more non-spatial strategies and showed allocentric-specific memory deficits. Furthermore, a detailed analysis of swimming strategies revealed allocentric memory deficits in the probe trial in female 3-month-old and male 7-month-old 5xFAD animals before the onset of severe reference memory deficits. Overall, we could demonstrate that spatial navigation deficits in 5xFAD mice are age- and sex-dependent, with female mice being more severely affected. In addition, the implementation of a search strategy classification system allowed an earlier detection of behavioral differences and therefore could be a powerful tool for preclinical drug testing in the 5xFAD model.

KUALITAS TEKNIK RENANG GAYA BEBAS ATLET TIRTA KALUANG

This study plans to decide the nature of the free-form swimming method. This examination is a sort of unmistakable exploration. The populace in this study were all swimming competitors from Tirta Kaluang Padang, adding up to 13 individuals. This testing was completed utilizing an all out inspecting method, which was 13 individuals. This examination instrument is through a free-form swimming strategy test. The information investigation strategy utilized recurrence circulation examination procedure with rate computation. The consequences of the review are, (1) The degree of evaluation for the nature of the body position procedure with an adequate level. (2) The degree of dominance of the nature of the arm procedure with an adequate level. (3) The degree of dominance of the nature of the foot development method with an adequate level. (4) The degree of dominance of the nature of breathing strategies with an adequate level. (5) The degree of dominance of the nature of coordination methods with an adequate level. Subsequently, the general examination results from 13 examples of swimming competitors from Tirta Kaluang Padang, acquired that the nature of free-form swimming procedures is in the adequate classification.

Sea Turtles Employ Drag-Reducing Techniques to Conserve Energy

Sea turtles are recognised as one of the ocean’s most remarkable migratory species, accomplishing journeys that cover thousands of kilometres. This fact is even more extraordinary when considering sea turtles consume mostly low-energy foods. The biology of sea turtles dominates the literature; however, the swimming strategies they employ to achieve their migratory success from a biomechanical and hydrodynamic viewpoint is relatively unexplored. In past research, the sea turtle’s upstroke has been debated among researchers as to whether it is passive or for thrust production. In this work, we recreate a model based on the green sea turtle (Chelonia mydas) and develop an ad hoc testing rig to uncover the secrets behind the sea turtle’s upstroke. Our findings suggest sea turtles utilise a passive upstroke that can substantially lower the animal’s drag coefficient to levels that cause insignificant losses in swim speed despite not developing any thrust force. This can conceivably save the animal a notable amount of energy as the upstroke is responsible for a large percentage of the overall limb beat cycle. These findings could potentially pave a path towards developing high-efficiency bioinspired underwater drone technologies.

Model-Based Design Optimization of Underwater Flagellate Propellers

A new family of soft underwater propellers has been recently presented. Mimicking the swimming strategy of bacterial flagella, these modules passively adapt to the surrounding fluid to provide a propulsive thrust. In the present paper we aim at further investigating the behaviour of this device and we address the optimization of its design towards improved swimming capabilities. This process is allowed by an accurate, yet simple, theoretical model which is able to precisely describe the robot's behaviour. The optimal prototype is fabricated, employing a composite material that is ad-hoc designed to provide the optimal stiffness. Finally, a simple robotic prototype is built and tested to validate the improved performances.

Ablation of the Presynaptic Protein Mover Impairs Learning Performance and Decreases Anxiety Behavior in Mice.

The presynaptic protein Mover/TPRGL/SVAP30 is absent in Drosophila and C. elegans and differentially expressed in synapses in the rodent brain, suggesting that it confers specific functions to subtypes of presynaptic terminals. In order to investigate how the absence of this protein affects behavior and learning, Mover knockout mice (KO) were subjected to a series of established learning tests. To determine possible behavioral and cognitive alterations, male and female 8-week-old KO and C57Bl/6J wildtype (WT) control mice were tested in a battery of memory and anxiety tests. Testing included the cross maze, novel object recognition test (NOR), the Morris water maze (MWM), the elevated plus maze (EPM), and the open field test (OF). Mover KO mice showed impaired recognition memory in the NOR test, and decreased anxiety behavior in the OF and the EPM. Mover KO did not lead to changes in working memory in the cross maze or spatial reference memory in the MWM. However, a detailed analysis of the swimming strategies demonstrated allocentric-specific memory deficits in male KO mice. Our data indicate that Mover appears to control synaptic properties associated with specific forms of memory formation and behavior, suggesting that it has a modulatory role in synaptic transmission.

Squirmer locomotion in a yield stress fluid

An axisymmetric squirmer in a Bingham viscoplastic fluid is studied numerically to determine the effect of a yield stress environment on locomotion. The nonlinearity of the governing equations necessitates numerical methods, which are accomplished by solving a variable-viscosity Stokes equation with a finite element approach. The effects of stroke modes, both pure and combined, are investigated, and it is found that for the treadmill or ‘neutral’ mode, the swimmer in a yield stress fluid has a lower swimming velocity and uses more power. However, the efficiency of swimming reaches its maximum at a finite yield limit. In addition, for higher yield limits, higher stroke modes can increase the swimming velocity and hydrodynamic efficiency of the treadmill swimmer. The higher-order odd-numbered squirming modes, particularly the third stroke mode, can generate propulsion by themselves that increases in strength as the viscoplastic nonlinearity increases to a specific limit. These results are closely correlated with the confinement effects induced by the viscoplastic rigid surface surrounding the swimming body, showing that swimmers in viscoplastic environments, both biological and artificial, could potentially employ other non-standard swimming strategies to optimize their locomotion.