A solution strategy to include the opening of the opercular slits in moving-mesh CFD models of suction feeding

The gill cover of fish and pre-metamorphic salamanders has a key role in suction feeding by acting as a one-way valve. It initially closes to avoid an inflow of water through the gill slits, after which it opens to allow outflow of the water that was sucked through the mouth into the expanded buccopharyngeal cavity. However, due to the inability of analytical models (relying on the continuity principle) to calculate a fluid flow through a shape-and-size-changing cavity with two openings, stringent boundary conditions had to be used in previously developed mathematical models after the moment of valve opening. By solving additionally for momentum conservation, computational fluid dynamics (CFD) has the capacity to dynamically simulate these flows, but this technique also faces complications to model a transition from closed to open valves. Here, I present a relatively simple solution strategy to incorporate valve opening, exemplified in an axisymmetrical model of a suction-feeding sunfish in ANSYS Fluent software. By controlling viscosity of a separately defined fluid entity at the opercular cavity region, early inflow can be blocked (high viscosity assigned) and later outflow can be allowed (changing viscosity to that of water). Finally, by analysing the CFD solution obtained for the sunfish model, a few new insights in the biomechanics of suction feeding will be discussed

New insights into muscle function during pivot feeding in seahorses

Seahorses, pipefish and their syngnathiform relatives are considered unique amongst fishes in using elastic recoil of post-cranial tendons to pivot the head extremely quickly towards small crustacean prey. It is known that pipefish activate the epaxial muscles for a considerable time before striking, at which rotations of the head and the hyoid are temporarily prevented to allow energy storage in the epaxial tendons. Here, we studied the motor control of this system in seahorses using electromyographic recordings of the epaxial muscles and the sternohyoideus-hypaxial muscles with simultaneous high-speed video recordings of prey capture. In addition we present the results from a stimulation experiment including the muscle hypothesised to be responsible for the locking and triggering of pivot feeding in seahorses (m. adductor arcus palatini). Our data confirmed that the epaxial pre-activation pattern observed previously for pipefish also occurs in seahorses. Similar to the epaxials, the sternohyoideus-hypaxial muscle complex shows prolonged anticipatory activity. Although a considerable variation in displacements of the mouth via head rotation could be observed, it could not be demonstrated that seahorses have control over strike distance. In addition, we could not identify the source of the kinematic variability in the activation patterns of the associated muscles. Finally, the stimulation experiment supported the previously hypothesized role of the m. adductor arcus palatini as the trigger in this elastic recoil system. Our results show that pre-stressing of both the head elevators and the hyoid retractors is taking place. As pre-activation of the main muscles involved in pivot feeding has now been demonstrated for both seahorses and pipefish, this is probably a generalized trait of Syngnathidae

Environment-dependent prey capture in the Atlantic mudskipper (Periophthalmus barbarus)

Few vertebrates capture prey in both the aquatic and the terrestrial environment due to the conflicting biophysical demands of feeding in water versus air. The Atlantic mudskipper (Periophthalmus barbarus) is known to be proficient at feeding in the terrestrial environment and feeds predominately in this environment. Given the considerable forward flow of water observed during the mouth-opening phase to assist with feeding on land, the mudskipper must alter the function of its feeding system to feed successfully in water. Here, we quantify the aquatic prey-capture kinematics of the mudskipper and compare this with the previously described pattern of terrestrial feeding. Prior to feeding in the aquatic environment, the gill slits open, allowing water to be expelled through the gill slits. The opposite happens in terrestrial feeding during which the gill slits remain closed at this point. In water, the expansive movements of the head are larger, amounting to a larger volume increase and are initiated slightly later than in the terrestrial environment. This implies the generation of strong suction flows when feeding in water. Consequently, the kinematic patterns of the hydrodynamic tongue during terrestrial feeding and aquatic suction feeding are similar, except for the amplitude of the volume increase and the active closing of the gill slits early during the terrestrial feeding strike. The mudskipper thus exhibits the capacity to change the kinematics of its feeding apparatus to enable successful prey capture in two disparate environments

Phenotypic flexibility of gape anatomy fine-tunes the aquatic prey-capture system of newts

A unique example of phenotypic flexibility of the oral apparatus is present in newts (Salamandridae) that seasonally change between an aquatic and a terrestrial habitat. Newts grow flaps of skin between their upper and lower jaws, the labial lobes, to partly close the corners of the mouth when they adopt an aquatic lifestyle during their breeding season. Using hydrodynamic simulations based on mu CT-scans and cranial kinematics during prey-capture in the smooth newt (Lissotriton vulgaris), we showed that this phenotypic flexibility is an adaptive solution to improve aquatic feeding performance: both suction distance and suction force increase by approximately 15% due to the labial lobes. As the subsequent freeing of the corners of the mouth by resorption of the labial lobes is assumed beneficial for the terrestrial capture of prey by the tongue, this flexibility of the mouth fine-tunes the process of capturing prey throughout the seasonal switching between water and land

An adaptive explanation for the horse-like shape of seahorses

The body shape of seahorses resembles the head and neck of horses because of their curved trunk, their ventrally bent head and their long snout. Seahorses evolved from ancestral, pipefish-like species, which have a straight body. Here, we use a biomechanical analysis and show that the seahorse's peculiar head, neck and trunk posture allows for the capture of small shrimps at larger distances from the eyes compared with pipefish. The results from the mathematical modelling were confirmed by kinematic data of prey-capturing syngnathids: compared with straight-bodied pipefish, all seahorse species studied consistently show an additional forward-reaching component in the path travelled by the mouth during their strikes at prey. This increased strike distance enlarges the volume of water they can probe for food, which is especially useful for tail-attached, sit-and-wait predators like seahorses. The biomechanics of prey capture thus provides a putative selective advantage that may explain the bending of the trunk into a horse-like shape

Kinematics of chisel-tooth digging by African mole-rats

Mole-rats are known to use their protruding, chisel-like incisors to dig underground networks of tunnels, but it remains unknown how these incisors are used to break and displace the soil. Theoretically, different excavation strategies can be used. Mole-rats could either use their head depressor muscles to power scooping motions of the upper incisors (by nose-down head rotations) or the lower incisors (by nose-up head rotations), or their jaw adductors to grab and break the soil after penetrating both sets of incisors into the ground, or a combination of these mechanisms. To identify how chisel-tooth digging works, a kinematic analysis of this behaviour was performed based on high-speed videos of 19 individuals from the African molerat species Fukomys micklemi placed inside transparent tubes in a laboratory setting. Our analysis showed that the soil is penetrated by both the upper and lower incisors at a relatively high gape angle, generally with the head rotated nose-up. Initially, the upper incisors remain approximately stationary to function as an anchor to allow an upward movement of the lower incisors to grab the soil. Next, a quick, nose-down rotation of the head further detaches the soil and drops the soil below the head. Consequently, both jaw adduction and head depression are jointly used to power tooth-digging in F. micklemi. The same mechanism, but with longer digging cycles, and soil being thrown down at smaller gape sizes, was used when digging in harder soil

Modulating yaw with an unstable rigid body and a course-stabilizing or steering caudal fin in the yellow boxfish (Ostracion cubicus)

Despite that boxfishes have a rigid carapace that restricts body undulation, they are highly manoeuvrable and manage to swim with remarkably dynamic stability. Recent research has indicated that the rigid body shape of boxfishes shows an inherently unstable response in its rotations caused by course-disturbing flows. Hence, any net stabilizing effect should come from the fishes' fins. The aim of the current study was to determine the effect of the surface area and orientation of the caudal fin on the yaw torque exerted on the yellow boxfish, Ostracion cubicus, a square cross-sectional shaped species of boxfish. Yaw torques quantified in a flow tank using a physical model with an attachable closed or open caudal fin at different body and tail angles and at different water flow speeds showed that the caudal fin is crucial for controlling yaw. These flow tank results were confirmed by computational fluid dynamics simulations. The caudal fin acts as both a course-stabilizer and rudder for the naturally unstable rigid body with regard to yaw. Boxfishes seem to use the interaction of the unstable body and active changes in the shape and orientation of the caudal fin to modulate manoeuvrability and stability

Modulation of yaw by the caudal fin in the yellow boxfish (<i>Ostracion cubicus</i>)

Boxfishes (Ostraciidae; Tetraodontiformes) have a rigid carapace which restricts body undulation. Swimming movements can only be generated by the fins which protrude from the carapace. Nevertheless, these fishes are highly manoeuvrable and manage to swim with remarkably dynamic stability. However, the rigid carapace of boxfishes shows an inherently unstable response in yaw caused by course-disturbing flows. Hence, any net stabilising effect should come from the fishes’ fins. Here, we aim to determine the effect of the surface area and orientation of the caudal fin on the yaw torque exerted on the square cross-sectional shaped yellow boxfish (Ostracion cubicus). Yaw torques were quantified in a flow tank using a 3D printed physical yellow boxfish model with an attachable closed or open caudal fin. The model was positioned at different body and tail angles and exposed to different water flow speeds. We show that the caudal fin is crucial for yaw control. These flow tank results were confirmed by computational fluid dynamics simulations. The caudal fin acts as both a course-stabiliser and rudder for the naturally unstable rigid carapace with regard to yaw. By using physical models and computer simulations, we quantitatively show that actively changing the shape and orientation of the caudal fin plays an important role in controlling yaw torque in yellow boxfish. Further study is needed to unravel how all components of the boxfishes’ locomotor apparatus function together, from a dynamic perspective, during lateral gust flows and turning