Allometric Scaling Reveals Evolutionary Constraint on Odonata Wing Cellularity via Critical Crack Length.

Scaling in insect wings is a complex phenomenon that seems pivotal in maintaining wing functionality. In this study, the relationship between wing size and the size, location, and shape of wing cells in dragonflies and damselflies (Odonata) is investigated, aiming to address the question of how these factors are interconnected. To this end, WingGram, the recently developed computer-vision-based software, is used to extract the geometric features of wing cells of 389 dragonflies and damselfly wings from 197 species and 16 families. It has been found that the cell length of the wings does not depend on the wing size. Despite the wide variation in wing length (8.42 to 56.5 mm) and cell length (0.1 to 8.5 mm), over 80% of the cells had a length ranging from 0.5 to 1.5 mm, which was previously identified as the critical crack length of the membrane of locust wings. An isometric scaling of cells is also observed with maximum size in each wing, which increased as the size increased. Smaller cells tended to be more circular than larger cells. The results have implications for bio-mimetics, inspiring new materials and designs for artificial wings with potential applications in aerospace engineering and robotics.

Optimization of Enzymatic and Chemical Decellularization of Native Porcine Heart Valves for the Generation of Decellularized Xenografts.

One of the most important medical interventions for individuals with heart valvular disease is heart valve replacement, which is not without substantial challenges, particularly for pediatric patients. Due to their biological properties and biocompatibility, natural tissue-originated scaffolds derived from human or animal sources are one type of scaffold that is widely used in tissue engineering. However, they are known for their high potential for immunogenicity. Being free of cells and genetic material, decellularized xenografts, consequently, have low immunogenicity and, thus, are expected to be tolerated by the recipient's immune system. The scaffold ultrastructure and ECM composition can be affected by cell removal agents. Therefore, applying an appropriate method that preserves intact the structure of the ECM plays a critical role in the final result. So far, there has not been an effective decellularization technique that preserves the integrity of the heart valve's ultrastructure while securing the least amount of genetic material left. This study demonstrates a new protocol with untraceable cells and residual DNA, thereby maximally reducing any chance of immunogenicity. The mechanical and biochemical properties of the ECM resemble those of native heart valves. Results from this study strongly indicate that different critical factors, such as ionic detergent omission, the substitution of Triton X-100 with Tergitol, and using a lower concentration of trypsin and a higher concentration of DNase and RNase, play a significant role in maintaining intact the ultrastructure and function of the ECM.

Mandible elemental composition and mechanical properties from distinct castes of the leafcutter ant Atta laevigata (Attini; Formicidae).

Leafcutter ant colonies are divided into castes with the individuals performing different tasks, based mostly on size. With the mandibles, the small minims care for the brood or the fungus, whereas the larger minors and mediae cut and transport plant material, with the ant size positively related to the material size. The mechanical properties and composition of the mandible cuticle have been previously tested in the soldiers as the largest caste, revealing that the cutting edges contained high contents of the cross-linking transition metal zinc (Zn). With regard to the smaller castes, no data are present. To study how the mandible size and function relates to its mechanical properties, we here tested the mandibles of minims, minors and mediae by nanoindentation. We found that the hardness (H) and Young's modulus (E) values increased with increasing ant size and that the mandible cutting edges in each caste have the highest H- and E-values. To gain insight into the origins of these properties, we characterized the elemental composition by energy-dispersive X-ray analysis, revealing that minors and mediae possessed higher content of Zn in the cutting edges in contrast to the minims containing significantly less Zn. This shows, that Zn content relates to higher mechanical property values. Additionally, it shows that all of these parameters can differ within a single species.

Tanning of the tarsal and mandibular cuticle in adult Anax imperator (Insecta: Odonata) during the emergence sequence.

The arthropod cuticle offers strength, protection, and lightweight. Due to its limit in expandability, arthropods have to moult periodically to grow. While moulting is beneficial in terms of parasite or toxin control, growth and adaptation to environmental conditions, it costs energy and leaves the soft animal's body vulnerable to injuries and desiccation directly after ecdysis. To investigate the temporal change in sclerotization and pigmentation during and after ecdysis, we combined macrophotography, confocal laser scanning microscopy, scanning electron microscopy and histological sectioning. We analysed the tarsal and mandibular cuticle of the blue emperor dragonfly to compare the progress of tanning for structures that are functionally involved during emergence (tarsus/tarsal claws) with structures whose functionality is required much later (mandibles). Our results show that: (i) the tanning of the tarsal and mandibular cuticle increases during emergence; (ii) the tarsal cuticle tans faster than the mandibular cuticle; (iii) the mandibles tan faster on the aboral than on the oral side; and (iv) both the exo- and the endocuticle are tanned. The change in the cuticle composition of the tarsal and mandibular cuticle reflects the demand for higher mechanical stability of these body parts when holding on to the substrate during emergence and during first walking or hunting attempts.

Mechanical and elemental characterization of ant mandibles: consequences for bite mechanics.

Mandible morphology has an essential role in biting performance, but the mandible cuticle can have regional differences in its mechanical properties. The effects of such a heterogeneous distribution of cuticle material properties in the mandible responses to biting loading are still poorly explored in chewing insects. Here, we tested the mechanical properties of mandibles of the ant species Formica cunicularia by nanoindentation and investigated the effects of the cuticular variation in Young's modulus (E) under bite loading with finite-element analysis (FEA). The masticatory margin of the mandible, which interacts with the food, was the hardest and stiffest region. To unravel the origins of the mechanical property gradients, we characterized the elemental composition by energy-dispersive X-ray spectroscopy. The masticatory margin possessed high proportions of Cu and Zn. When incorporated into the FEA, variation in E effectively changed mandible stress patterns, leading to a relatively higher concentration of stresses in the stiffer mandibular regions and leaving the softer mandible blade with relatively lower stress. Our results demonstrated the relevance of cuticle E heterogeneity in mandibles under bite loading, suggesting that the accumulation of transition metals such as Cu and Zn has a relevant correlation with the mechanical characteristics in F. cunicularia mandibles.

Wear patterns of radular teeth in Loligo vulgaris (Cephalopoda; Mollusca) are related to their structure and mechanical properties.

Radular teeth have to cope with wear, when interacting with ingesta. In some molluscan taxa, wear-coping mechanisms, related to the incorporation of high contents of iron or silica, have been previously determined. For most species, particularly for those which possess radulae without such incorporations, wear-coping mechanisms are understudied. In the present study, we documented and characterized the wear on radular teeth in the model species Loligo vulgaris (Cephalopoda). By applying a range of methods, the elementary composition and mechanical properties of the teeth were described, to gain insight into mechanisms for coping with abrasion. It was found that the tooth regions that are prone to wear are harder and stiffer. Additionally, the surfaces interacting with the ingesta possessed a thin coating with high contents of silicon, probably reducing abrasion. The here presented data may serve as an example of systematic study of radular wear, in order to understand the relationship between the structure of radular teeth and their properties.

Insect attachment on waxy plant surfaces: the effect of pad contamination by different waxes.

This study focuses on experimental testing of the contamination hypothesis and examines how the contamination of insect adhesive pads with three-dimensional epicuticular waxes of different plant species contributes to the reduction of insect attachment. We measured traction forces of tethered Chrysolina fastuosa male beetles having hairy adhesive pads on nine wax-bearing plant surfaces differing in both shape and dimensions of the wax structures and examined insect adhesive organs after they have contacted waxy substrates. For comparison, we performed the experiments with the same beetle individuals on a clean glass sample just before (gl1) and immediately after (gl2) the test on a plant surface. The tested insects showed a strong reduction of the maximum traction force on all waxy plant surfaces compared to the reference experiment on glass (gl1). After beetles have walked on waxy plant substrates, their adhesive pads were contaminated with wax material, however, to different extents depending on the plant species. The insects demonstrated significantly lower values of both the maximum traction force and the first peak of the traction force and needed significantly longer time to reach the maximum force value in the gl2 test than in the gl1 test. These effects were especially pronounced in cases of the plant surfaces covered with wax projections having higher aspect ratios. The data obtained clearly indicated the impact of waxy plant surfaces on the insect ability to subsequently attach to the clean smooth surface. This effect is caused by the contamination of adhesive pads and experimentally supports the contamination hypothesis.

Streptomyces profundus sp. nov., a novel marine actinobacterium isolated from deep-sea sediment of Madeira Archipelago, Portugal.

A novel strain, MA3_2.13T, was isolated from deep-sea sediment of Madeira Archipelago, Portugal, and characterized using a polyphasic approach. This strain produced dark brown soluble pigments, bronwish black substrate mycelia and an aerial mycelium with yellowish white spores, when grown on GYM 50SW agar. The main respiratory quinones were MK-10(H4), MK-10(H6) and MK-10(H8). Diphosphatidylglycerol, phosphatidylethanolamine, three unidentified phospholipids and two glycophospholipids were identified as the main phospholipids. The major cellular fatty acids were iso-C16 : 1, iso-C16 : 0, anteiso-C17 : 1 and anteiso-C17 : 0. Phylogenetic analyses based on 16S rRNA gene showed that strain MA3_2.13T is a member of the genus Streptomyces and was most closely related to Streptomyces triticirhizae NEAU-YY642T (NR_180032.1; 16S rRNA gene similarity 97.9 %), Streptomyces sedi YIM 65188T (NR_044582.1; 16S rRNA gene similarity 97.4 %), Streptomyces mimosae 3MP-10T (NR_170412.1; 16S rRNA gene similarity 97.3 %) and Streptomyces zhaozhouensis NEAU-LZS-5T (NR_133874.1; 16S rRNA gene similarity 97.0 %). Genome pairwise comparisons with closest related type strains retrieved values below the threshold for species delineation suggesting that strain MA3_2.13T represents a new branch within the genus Streptomyces. Based on these results, strain MA3_2.13T (=DSM 115980T=LMG 33094T) is proposed as the type strain of a novel species of the genus Streptomyces, for which the name Streptomyces profundus sp. nov. is proposed.

Multiple forces facilitate the aquatic acrobatics of grasshopper and bioinspired robot.

Aquatic locomotion is challenging for land-dwelling creatures because of the high degree of fluidity with which the water yields to loads. We surprisingly found that the Chinese rice grasshopper Oxya chinensis, known for its terrestrial acrobatics, could swiftly launch itself off the water's surface in around 25 ms and seamlessly transition into flight. Biological observations showed that jumping grasshoppers use their front and middle legs to tilt up bodies first and then lift off by propelling the water toward the lower back with hind legs at angular speeds of up to 18°/ms, whereas the swimming grasshoppers swing their front and middle legs in nearly horizontal planes and move hind legs less violently (~8°/ms). Force measurement and model analysis indicated that the weight support could be achieved by hydrostatics which are proportionate to the mass of the grasshoppers, while the propulsions for motion are derived from the controlled limb-water interactions (i.e., the hydrodynamics). After learning the structural and behavioral strategies of the grasshoppers, a robot was created and was capable of swimming and jumping on the water surface like the insects, further demonstrating the effectiveness of decoupling the challenges of aquatic locomotion by the combined use of the static and dynamic hydro forces. This work not only uncovered the combined mechanisms responsible for facilitating aquatic acrobatics in this species but also laid a foundation for developing bioinspired robots that can locomote across multiple media.

TriTrap: A Robotic Gripper Inspired by Insect Tarsal Chains.

Gripping, holding, and moving objects are among the main functional purposes of robots. Ever since automation first took hold in society, optimizing these functions has been of high priority, and a multitude of approaches has been taken to enable cheaper, more reliable, and more versatile gripping. Attempts are ongoing to reduce grippers' weight, energy consumption, and production and maintenance costs while simultaneously improving their reliability, the range of eligible objects, working loads, and environmental independence. While the upper bounds of precision and flexibility have been pushed to an impressive level, the corresponding solutions are often dependent on support systems (e.g., sophisticated sensors and complex actuation machinery), advanced control paradigms (e.g., artificial intelligence and machine learning), and typically require more maintenance owed to their complexity, also increasing their cost. These factors make them unsuited for more modest applications, where moderate to semi-high performance is desired, but simplicity is required. In this paper, we attempt to highlight the potential of the tarsal chain principle on the example of a prototype biomimetic gripping device called the TriTrap gripper, inspired by the eponymous tarsal chain of insects. Insects possess a rigid exoskeleton that receives mobility due to several joints and internally attaching muscles. The tarsus (foot) itself does not contain any major intrinsic muscles but is moved by an extrinsically pulled tendon. Just like its biological counterpart, the TriTrap gripping device utilizes strongly underactuated digits that perform their function using morphological encoding and passive conformation, resulting in a gripper that is versatile, robust, and low cost. Its gripping performance was tested on a variety of everyday objects, each of which represented different size, weight, and shape categories. The TriTrap gripper was able to securely hold most of the tested objects in place while they were lifted, rotated, and transported without further optimization. These results show that the insect tarsus selected approach is viable and warrants further development, particularly in the direction of interface optimization. As such, the main goal of the TriTrap gripper, which was to showcase the tarsal chain principle as a viable approach to gripping in general, was achieved.

Mandible mechanical properties and composition of the larval Glossosoma boltoni (Trichoptera, Insecta).

Insect feeding structures, such as mandibles, interact with the ingesta (food or/and substrate) and can be adapted in morphology, composition of material and mechanical properties. The foraging on abrasive ingesta, as on algae covering rocks, is particularly challenging because the mandibles will be prone to wear and structural failure, thus suggesting the presence of mandibular adaptations to accompany this feeding behavior. Adaptations to this are well studied in the mouthparts of molluscs and sea urchins, but for insects there are large gaps in our knowledge. In this study, we investigated the mandibles of a grazing insect, the larvae of the trichopteran Glossosoma boltoni. Using scanning electron microscopy, wear was documented on the mandibles. The highest degree was identified on the medial surface of the sharp mandible tip. Using nanoindentation, the mechanical properties, such as hardness and Young's modulus, of the medial and lateral mandible cuticles were tested. We found, that the medial cuticle of the tip was significantly softer and more flexible than the lateral one. These findings indicate that a self-sharpening mechanism is present in the mandibles of this species, since the softer medial cuticle is probably abraded faster than the harder lateral one, leading to sharp mandible tips. To investigate the origins of these properties, we visualized the degree of tanning by confocal laser scanning microscopy. The autofluorescence signal related to the mechanical property gradients. The presence of transition and alkaline earth metals by energy dispersive X-ray spectroscopy was also tested. We found Ca, Cl, Cu, Fe, K, Mg, Mn, P, S, Si, and Zn in the cuticle, but the content was very low and did not correlate with the mechanical property values.

Attachment performance of the ectoparasitic seal louse Echinophthirius horridus.

Marine mammals host a great variety of parasites, which usually co-evolved in evolutionary arms races. However, little is known about the biology of marine mammal insect parasites, and even less about physical aspects of their life in such a challenging environment. One of 13 insect species that manage to endure long diving periods in the open sea is the seal louse, Echinophthirius horridus, parasitising true seals. Its survival depends on its specialised adaptations for enduring extreme conditions such as hypoxia, temperature changes, hydrostatic pressure, and strong drag forces during host dives. To maintain a grip on the seal fur, the louse's leg morphology is equipped with modified snap hook claws and soft pad-like structures that enhance friction. Through techniques including CLSM, SEM, and histological staining, we have examined the attachment system's detailed structure. Remarkably, the seal louse achieves exceptional attachment forces on seal fur, with safety factors (force per body weight) reaching 4500 in average measurements and up to 18000 in peak values, indicating superior attachment performance compared to other insect attachment systems. These findings underscore the louse's remarkable adaptations for life in a challenging marine environment, shedding light on the relationship between structure and function in extreme ecological niches.

Heartworm and seal louse: Trends in prevalence, characterisation of impact and transmission pathways in a unique parasite assembly on seals in the North and Baltic Sea.

The ectoparasitic seal louse, Echinophthirius horridus infects harbour (Phoca vitulina) and grey seals (Halichoerus grypus) in the North and Baltic Sea. The endoparasitic heartworm Acanthocheilonema spirocauda parasitizes the right heart and blood vessels of harbour seals. The complete lifecycle of the heartworm is not entirely understood although the seal louse is assumed to serve as vector for its transmission. Knowledge about the impact of both parasite species on host health are scarce. In this study, necropsy data and archived parasites of harbour and grey seals in German waters were analysed to determine long-term seal louse (SLP) and heartworm prevalence (HWP) from 2014 to 2021. Histology, microbiology and scanning electron microscopy (SEM) were applied on seal louse infected and uninfected skin to investigate associated lesions and the health impact. During the study period, HWP in harbour seals was 13%, the SLP in harbour seals was 4% and in grey seals 10%. HWP of harbour seals was significantly higher during the winter months compared to the summer. SLP in adults was significantly higher in comparison to juvenile harbour seals. SLP varied significantly between grey seals from the North and Baltic Sea. Filarial nematodes were detected in the haemocoel, pharynx, and intestine of E. horridus highlighting the seal louse as vector for heartworms. Alopecia and folliculitis were associated with the attachment posture of E. horridus and microbiological investigations isolated bacteria commonly associated with folliculitis.

The effect of surface topography on the ball-rolling ability of Kheper lamarcki (Scarabaeidae).

The most effective way to avoid intense inter- and intra-specific competition at the dung source, and to increase the distance to the other competitors, is to follow a single straight bearing. While ball-rolling dung beetles manage to roll their dung balls along nearly perfect straight paths when traversing flat terrain, the paths that they take when traversing more complex (natural) terrain are not well understood. In this study, we investigate the effect of complex surface topographies on the ball-rolling ability of Kheper lamarcki. Our results reveal that ball-rolling trajectories are strongly influenced by the characteristic scale of the surface structure. Surfaces with an increasing similarity between the average distance of elevations and the ball radius cause progressively more difficulties during ball transportation. The most important factor causing difficulties in ball transportation appears to be the slope of the substrate. Our results show that, on surfaces with a slope of 7.5 deg, more than 60% of the dung beetles lose control of their ball. Although dung beetles still successfully roll their dung ball against the slope on such inclinations, their ability to roll the dung ball sideways diminishes. However, dung beetles do not seem to adapt their path on inclines such that they roll their ball in the direction against the slope. We conclude that dung beetles strive for a straight trajectory away from the dung pile, and that their actual path is the result of adaptations to particular surface topographies.

Attachment discs of the diving bell spider Argyroneta aquatica.

To adhere their silk threads for the construction of webs and to fix the dragline, spiders produce attachment discs of piriform silk. Uniquely, the aquatic spider Argyroneta aquatica spends its entire life cycle underwater. Therefore, it has to glue its attachment discs to substrates underwater. Here we show that Argyroneta aquatica applies its thread anchors within an air layer around the spinnerets maintained by superhydrophobic setae. During spinning, symmetric movements of the spinnerets ensure retaining air in the contact area. The flat structure of the attachment discs is thought to facilitate fast curing of the piriform adhesive cement and improves the resistance against drag forces. Pull-off tests on draglines connected with attachment discs on different hydrophilic substrates point to dragline rupture as the failure mode. The Young´s modulus of the dragline (8.3 GPa) is within the range as in terrestrial spiders. The shown structural and behavioral adaptations can be the model for new artificial underwater gluing devices.

Holding in the stream: convergent evolution of suckermouth structures in Loricariidae (Siluriformes).

Suckermouth armoured catfish (Loricariidae) are a highly speciose and diverse freshwater fish family, which bear upper and lower lips forming an oral disc. Its hierarchical organisation allows the attachment to various natural surfaces. The discs can possess papillae of different shapes, which are supplemented, in many taxa, by small horny projections, i.e. unculi. Although these attachment structures and their working mechanisms, which include adhesion and interlocking, are rather well investigated in some selected species, the loricariid oral disc is unfortunately understudied in the majority of species, especially with regard to comparative aspects of the diverse oral structures and their relationship to the ecology of different species. In the present paper, we investigated the papilla and unculi morphologies in 67 loricariid species, which inhabit different currents and substrates. We determined four papilla types and eight unculi types differing by forms and sizes. Ancestral state reconstructions strongly suggest convergent evolution of traits. There is no obvious correlation between habitat shifts and the evolution of specific character states. From handling the structures and from drying artefacts we could infer some information about their material properties. This, together with their shape, enabled us to carefully propose hypotheses about mechanisms of interactions of oral disc structures with natural substrates typical for respective fish species.

Anti-Adhesive Surfaces Inspired by Bee Mandible Surfaces.

Propolis, a naturally sticky substance used by bees to secure their hives and protect the colony from pathogens, presents a fascinating challenge. Despite its adhesive nature, honeybees adeptly handle propolis with their mandibles. Previous research has shown a combination of an anti-adhesive fluid layer and scale-like microstructures on the inner surface of bee mandibles. Our aim was to deepen our understanding of how surface energy and microstructure influence the reduction in adhesion for challenging substances like propolis. To achieve this, we devised surfaces inspired by the intricate microstructure of bee mandibles, employing diverse techniques including roughening steel surfaces, creating lacquer structures using Bénard cells, and moulding resin surfaces with hexagonal patterns. These approaches generated patterns that mimicked the bee mandible structure to varying degrees. Subsequently, we assessed the adhesion of propolis on these bioinspired structured substrates. Our findings revealed that on rough steel and resin surfaces structured with hexagonal dimples, propolis adhesion was significantly reduced by over 40% compared to unstructured control surfaces. However, in the case of the lacquer surface patterned with Bénard cells, we did not observe a significant reduction in adhesion.

Producing and Testing Prototype Tissue-Engineered 3D Tri-Leaflet Valved Stents on Biodegradable Poly-ε-Caprolactone Scaffolds.

Transcatheter pulmonary valve replacement is a minimally-invasive alternative treatment for right ventricular outflow tract dysfunction and has been rapidly evolving over the past years. Heart valve prostheses currently available still have major limitations. Therefore, one of the significant challenges for the future is the roll out of transcatheter tissue engineered pulmonary valve replacement to more patients. In the present study, biodegradable poly-ε-caprolactone (PCL) nanofiber scaffolds in the form of a 3D leaflet matrix were successfully seeded with human endothelial colony-forming cells (ECFCs), human induced pluripotent stem cell-derived MSCs (hMSCs), and porcine MSCs (pMSCs) for three weeks for the generation of 3D tissue-engineered tri-leaflet valved stent grafts. The cell adhesion, proliferation, and distribution of these 3D heart leaflets was analyzed using fluorescence microscopy and scanning electron microscopy (SEM). All cell lineages were able to increase the overgrown leaflet area within the three-week timeframe. While hMSCs showed a consistent growth rate over the course of three weeks, ECFSs showed almost no increase between days 7 and 14 until a growth spurt appeared between days 14 and 21. More than 90% of heart valve leaflets were covered with cells after the full three-week culturing cycle in nearly all leaflet areas, regardless of which cell type was used. This study shows that seeded biodegradable PCL nanofiber scaffolds incorporated in nitinol or biodegradable stents will offer a new therapeutic option in the future.

Particle binding capacity of snail saliva.

Gastropods forage with their radula, a thin chitinous membrane with embedded teeth, which scratch across the substrate to lose food particles. During this interaction, the risk of loosening particles is obvious without having a specialized mechanism holding them on the tooth surface. As mucus secretions are essential in molluscan life cycles and the locomotion and attachment gels are known to have an instant high adhesion, we have hypothesized that the saliva could support particle retention during feeding. As adhesion of snail saliva was not studied before, we present here an experimental setup to test its particle-binding capacity using a large land snail (Lissachatina fulica, Stylommatophora, Heterobranchia). This experiment was also applied to the gels produced by the snail foot for comparison and can be potentially applied to various fluids present at a small volume in the future. We found, that the saliva has high particle retention capacity that is comparable to the foot glue of the snail. To gain some insight into the properties of the saliva, we additionally studied it in the scanning electron microscope, estimated its viscosity in a de-wetting experiment, and investigated its elemental composition using energy dispersive X-ray spectroscopy reveling higher contents of Ca, Zn and other potential cross-linkers similar to those found in the glue.

Locomotory Behavior of Water Striders with Amputated Legs.

The stability of the body during locomotion is a fundamental requirement for walking animals. The mechanisms that coordinate leg movement patterns are even more complex at water-air interfaces. Water striders are agile creatures on the water surface, but they can be vulnerable to leg damage, which can impair their movement. One can assume the presence of certain compensatory biomechanical factors that are involved in the maintenance of postural balance lost after an amputation. Here, we studied changes in load distribution among the legs and assessed the effects of amputation on the locomotory behavior and postural defects that may increase the risk of locomotion failure. Apparently, amputees recover a stable posture by applying leg position modifications (e.g., widening the stance) and by load redistribution to the remaining legs. Water striders showed steering failure after amputation in all cases. Amputations affected locomotion by (1) altering motion features (e.g., shorter swing duration of midlegs), (2) functional constraints on legs, (3) shorter travelled distances, and (4) stronger deviations in the locomotion path. The legs functionally interact with each other, and removal of one leg has detrimental effects on the others. This research may assist the bioinspired design of aquatic robots.