Curvature-induced stiffening of a fish fin

How fish modulate their fin stiffness during locomotive manoeuvres remains unknown. We show that changing the fin's curvature modulates its stiffness. Modelling the fin as bendable bony rays held together by a membrane, we deduce that fin curvature is manifested as a misalignment of the principal bending axes between neighbouring rays. An external force causes neighbouring rays to bend and splay apart, and thus stretches the membrane. This coupling between bending the rays and stretching the membrane underlies the increase in stiffness. Using analysis of a 3D reconstruction of a Mackerel (Scomber japonicus) pectoral fin, we calculate the range of stiffnesses this fin is expected to span by changing curvature. The 3D reconstruction shows that, even in its geometrically flat state, a functional curvature is embedded within the fin microstructure owing to the morphology of individual rays. Since the ability of a propulsive surface to transmit force to the surrounding fluid is limited by its stiffness, the fin curvature controls the coupling between the fish and its surrounding fluid. Thereby, our results provide mechanical underpinnings and morphological predictions for the hypothesis that the spanned range of fin stiffnesses correlates with the behaviour and the ecological niche of the fish

Reynolds number limits for jet propulsion: A numerical study of simplified jellyfish

The Scallop Theorem states that reciprocal methods of locomotion, such as jet propulsion or paddling, will not work in Stokes flow (Reynolds number = 0). In nature the effective limit of jet propulsion is still in the range where inertial forces are significant. It appears that almost all animals that use jet propulsion swim at Reynolds numbers (Re) of about 5 or more. Juvenile squid and octopods hatch from the egg already swimming in this inertial regime. The limitations of jet propulsion at intermediate Re is explored here using the immersed boundary method to solve the two-dimensional Navier Stokes equations coupled to the motion of a simplified jellyfish. The contraction and expansion kinematics are prescribed, but the forward and backward swimming motions of the idealized jellyfish are emergent properties determined by the resulting fluid dynamics. Simulations are performed for both an oblate bell shape using a paddling mode of swimming and a prolate bell shape using jet propulsion. Average forward velocities and work put into the system are calculated for Reynolds numbers between 1 and 320. The results show that forward velocities rapidly decay with decreasing Re for all bell shapes when Re < 10. Similarly, the work required to generate the pulsing motion increases significantly for Re < 10. When compared actual organisms, the swimming velocities and vortex separation patterns for the model prolate agree with those observed in Nemopsis bachei. The forward swimming velocities of the model oblate jellyfish after two pulse cycles are comparable to those reported for Aurelia aurita, but discrepancies are observed in the vortex dynamics between when the 2D model oblate jellyfish and the organism

The Functional Implications of Anuran Metamorphosis for Survival, Locomotor Performance, and Limb Bone Mechanical Properties

Many organisms must contend with navigating their environments from birth. An organism could be classified as – and is often studied – in the context of locomotion through a single habitat type. However, many organisms must contend with a wide variety of environmental obstacles and substrates. What’s more, a large group of animals, Lissamphibia, do so while undergoing drastic transformation of their morphology and locomotor appendages. This transformation, term metamorphosis, typically coincides with a movement from water as a tadpole, to land as a frog or salamander. Many studies have associated this transitionary period with decreased locomotor performance and worse survival compared to amphibians pre or post metamorphosis and have termed this paradigm as the “Adaptive Peaks Hypothesis”. This body of work sought to address the generality of the Adaptive Peaks Hypothesis by studying the locomotor performance, development, and survival of a fully aquatic frog, Xenopus laevis. Swimming performance and survival of frogs across developmental stages were measured using high speed cameras and a predation tank where morphological traits were compared between (non)survivors post predation. Another subset of frogs, including aquatic X. laevis and more terrestrial L catesbianus, were used to quantify bone mineral density and material properties of developing frogs. Finally, the aquatic frog, Xenopus laevis, underwent jumping performance trials using high speed cameras and force plates to determine how well an aquatic frog could perform out of the water. The adaptive peaks hypothesis is not as robust as once thought. Xenopus laevis did not experience a performance decrease during metamorphosis, despite similar capture rates from the predators. Furthermore, the bone development of the femur differed from the more terrestrial frogs such that the less dense bones had greater stiffness in Xenopus. Finally, despite being fully aquatic, Xenopus frogs were still capable jumpers across two incline treatments with greater variability in jump performance in the more juvenile stages. Taken together, this body of work builds upon our understanding of animal locomotion by addressing how animals with complex life cycles navigate complex habitats

Chaotic exploration and learning of locomotion behaviours

We present a general and fully dynamic neural system, which exploits intrinsic chaotic dynamics, for the real-time goal-directed exploration and learning of the possible locomotion patterns of an articulated robot of an arbitrary morphology in an unknown environment. The controller is modeled as a network of neural oscillators that are initially coupled only through physical embodiment, and goal-directed exploration of coordinated motor patterns is achieved by chaotic search using adaptive bifurcation. The phase space of the indirectly coupled neural-body-environment system contains multiple transient or permanent self-organized dynamics, each of which is a candidate for a locomotion behavior. The adaptive bifurcation enables the system orbit to wander through various phase-coordinated states, using its intrinsic chaotic dynamics as a driving force, and stabilizes on to one of the states matching the given goal criteria. In order to improve the sustainability of useful transient patterns, sensory homeostasis has been introduced, which results in an increased diversity of motor outputs, thus achieving multiscale exploration. A rhythmic pattern discovered by this process is memorized and sustained by changing the wiring between initially disconnected oscillators using an adaptive synchronization method. Our results show that the novel neurorobotic system is able to create and learn multiple locomotion behaviors for a wide range of body configurations and physical environments and can readapt in realtime after sustaining damage

Understanding the agility of running birds: Sensorimotor and mechanical factors in avian bipedal locomotion

Birds are a diverse and agile lineage of vertebrates that all use bipedal locomotion for at least part of their life. Thus birds provide a valuable opportunity to investigate how biomechanics and sensorimotor control are integrated for agile bipedal locomotion. This review summarizes recent work using terrain perturbations to reveal neuromechanical control strategies used by ground birds to achieve robust, stable and agile running. Early experiments in running guinea fowl aimed to reveal the immediate intrinsic mechanical response to an unexpected drop ('pothole') in terrain. When navigating the pothole, guinea fowl experience large changes in leg posture in the perturbed step, which correlates strongly with leg loading and perturbation recovery. Analysis of simple theoretical models of running has further confirmed the crucial role of swing-leg trajectory control for regulating foot contact timing and leg loading in uneven terrain. Coupling between body and leg dynamics results in an inherent trade-off in swing leg retraction rate for fall avoidance versus injury avoidance. Fast leg retraction minimizes injury risk, but slow leg retraction minimizes fall risk. Subsequent experiments have investigated how birds optimize their control strategies depending on the type of perturbation (pothole, step, obstacle), visibility of terrain, and with ample practice negotiating terrain features. Birds use several control strategies consistently across terrain contexts: 1) independent control of leg angular cycling and leg length actuation, which facilitates dynamic stability through simple control mechanisms, 2) feedforward regulation of leg cycling rate, which tunes foot-contact timing to maintain consistent leg loading in uneven terrain (minimizing fall and injury risks), 3) load-dependent muscle actuation, which rapidly adjusts stance push-off and stabilizes body mechanical energy, and 4) multi-step recovery strategies that allow body dynamics to transiently vary while tightly regulating leg loading to minimize risks of fall and injury. In future work, it will be interesting to investigate the learning and adaptation processes that allow animals to adjust neuromechanical control mechanisms over short and long timescales