The exceptionally high speed at which syngnathid fishes are able to rotate their snout towards prey and capture it by suction is potentially caused by a catapult mechanism in which the energy previously stored in deformed elastic elements is suddenly released. According to this hypothesis, tension is built up in tendons of the post-cranial muscles before prey capture is initiated. Next, an abrupt elastic recoil generates high-speed dorsal rotation of the head and snout, rapidly bringing the mouth close to the prey, thus enabling the pipefish to be close enough to engulf the prey by suction. However, no experimental evidence exists for such a mechanism of mechanical power amplification during feeding in these fishes. To test this hypothesis, inverse dynamical modelling based upon kinematic data from high-speed videos of prey capture in bay pipefish Syngnathus leptorhynchus, as well as electromyography of the muscle responsible for head rotation (the epaxial muscle) was performed. The remarkably high instantaneous muscle-mass-specific power requirement calculated for the initial phase of head rotation (up to 5795 W kg K1 ), as well as the early onset times of epaxial muscle activity (often observed more than 300 ms before the first externally discernible prey capture motion), support the elastic power enhancement hypothesis.
Suction feeding is pervasive among aquatic vertebrates, and our understanding of the functional morphology and biomechanics of suction feeding has recently been advanced by combining experimental and modeling approaches. Key advances include the visualization of the patterns of flow in front of the mouth of a feeding fish, the measurement of pressure inside their mouth cavity, and the employment of analytical and computational models. Here, we review the key components of the morphology and kinematics of the suction-feeding system of anatomically generalized, adult rayfinned fishes, followed by an overview of the hydrodynamics involved. In the suction-feeding apparatus, a strong mechanistic link among morphology, kinematics, and the capture of prey is manifested through the hydrodynamic interactions between the suction flows and solid surfaces (the mouth cavity and the prey). It is therefore a powerful experimental system in which the ecology and evolution of the capture of prey can be studied based on first principals.
SUMMARY Scaling effects on the kinematics of suction feeding in fish remain poorly understood, at least partly because of the inconsistency of the results of the existing experimental studies. Suction feeding is mechanically distinct from most other type of movements in that negative pressure inside the buccal cavity is thought to be the most important speed-limiting factor during suction. However, how buccal pressure changes with size and how this influences the speed of buccal expansion is unknown. In this paper, the effects of changes in body size on kinematics of suction feeding are studied in the catfish Clarias gariepinus. Video recordings of prey-capturing C. gariepinus ranging in total length from 111 to 923 mm were made,from which maximal displacements, velocities and accelerations of several elements of the cranial system were determined. By modelling the observed expanding head of C. gariepinus as a series of expanding hollow elliptical cylinders, buccal pressure and power requirement for the expansive phase of prey capture were calculated for an ontogenetic sequence of catfish. We found that angular velocities decrease approximately proportional with increasing cranial size, while linear velocities remain more or less constant. Although a decreasing (angular) speed of buccal expansion with increasing size could be predicted (based on calculations of power requirement and the expected mass-proportional scaling of available muscular power in C. gariepinus), the observed drop in (angular) speed during growth exceeds these predictions. The calculated muscle-mass-specific power output decreases significantly with size, suggesting a relatively lower suction effort in the larger catfish compared with the smaller catfish.
Detailed morphological analyses have identified a number of different mechanical pathways by which the morphologically complex cranial system of fishes can achieve mouth opening and hyoid depression. However, many of these proposed mechanisms remain untested. Furthermore, very little is known about the precise timing of activity of each of these mechanisms, and about the magnitude of each mechanism's total contribution to its proposed function. In the present study, all mouth opening and hyoid depression mechanisms described for Clarias gariepinus, an air-breathing catfish, are analysed. Highspeed X-ray videos were recorded during prey capture of three catfish implanted with small, radio-opaque markers in the cranial elements potentially involved. A kinematic analysis was performed from which data were used as input in planar four-bar models. This analysis shows that the opercular mouth-opening mechanism initiates mouth opening, but is not able to cause the complete mouth openings as observed on the X-ray videos. The latter is accomplished through the protractor hyoidei muscles, which couple hyoid depression to lower jaw depression in a four-bar system and also reinforce lower jaw depression by shortening during the final stage of mouth opening. Although the angulo-ceratohyal ligament was previously hypothesised to play a part in mouth opening, our results show that it probably does not, but rather functions as a hyoid-elevator during mouth closure. Finally, hyoid depression is exclusively achieved by the four-bar mechanism involving neurocranial elevation and pectoral girdle retraction, generally without any reinforcement by shortening of the sternohyoideus muscle. In contrast to the results from a recent analysis on sunfish, the catfish's sternohyoideus gradually elongates during hyoid depression
The magnitude of sub-ambient pressure inside the bucco-pharyngeal cavity of aquatic animals is generally considered a valuable metric of suction feeding performance. However, these pressures do not provide a direct indication of the effect of the suction act on the movement of the prey item. Especially when comparing suction performance of animals with differences in the shape of the expanding bucco-pharyngeal cavity, the link between speed of expansion, water velocity, force exerted on the prey and intra-oral pressure remains obscure. By using mathematical models of the heads of catfishes, a morphologically diverse group of aquatic suction feeders, these relationships were tested. The kinematics of these models were fine-tuned to transport a given prey towards the mouth in the same way. Next, the calculated pressures inside these models were compared. The results show that no simple relationship exists between the amount of generated sub-ambient pressure and the force exerted on the prey during suction feeding, unless animals of the same species are compared. Therefore, for evaluating suction performance in aquatic animals in future studies, the focus should be on the flow velocities in front of the mouth, for which a direct relationship exists with the hydrodynamic force exerted on prey.
Despite repeated acquisitions of aquatic or semi‐aquatic lifestyles revolving around piscivory, snakes have not evolved suction feeding. Instead, snakes use frontally or laterally directed strikes to capture prey under water. If the aquatic medium constrains strike performance because of its physical properties, we predict morphological and functional convergence in snakes that use similar strike behaviours. Here we use natricine snakes to test for such patterns of convergence in morphology and function. Our data show that frontal strikers have converged on a similar morphology characterized by narrow elongate heads with a reduced projected frontal surface area. Moreover, simple computational fluid dynamics models show that the observed morphological differences are likely biologically relevant as they affect the flow of water around the head. In general, our data suggest that the direction of evolution may be predictable if constraints are strong and evolutionary solutions limited.
The shape of the carapace protecting the body of boxfishes has been attributed an important hydrodynamic role in drag reduction and in providing automatic, flow-direction realignment and is therefore used in bioinspired design of cars. However, tight swimming-course stabilization is paradoxical given the frequent, high-performance manoeuvring that boxfishes display in their spatially complex, coral reef territories. Here, by performing flow-tank measurements of hydrodynamic drag and yaw moments together with computational fluid dynamics simulations, we reverse several assumptions about the hydrodynamic role of the boxfish carapace. Firstly, despite serving as a model system in aerodynamic design, drag-reduction performance was relatively low compared with more generalized fish morphologies. Secondly, the current theory of course stabilization owing to flow over the boxfish carapace was rejected, as destabilizing moments were found consistently. This solves the boxfish swimming paradox: destabilizing moments enhance manoeuvrability, which is in accordance with the ecological demands for efficient turning and tilting.
During the evolutionary transition from fish to tetrapods, a shift from uni- to bidirectional suction feeding systems followed a reduction in the gill apparatus. Such a shift can still be observed during metamorphosis of salamanders, although many adult salamanders retain their aquatic lifestyle and feed by high-performance suction. Unfortunately, little is known about the interplay between jaws and hyobranchial motions to generate bidirectional suction flows. Here, we study the cranial morphology, as well as kinematic and hydrodynamic aspects related to prey capture in the Chinese giant salamander (Andrias davidianus). Compared with fish and previously studied amphibians, A. davidianus uses an alternative suction mechanism that mainly relies on accelerating water by separating the ‘plates’ formed by the long and broad upper and lower jaw surfaces. Computational fluid dynamics simulations, based on three-dimensional morphology and kinematical data from high-speed videos, indicate that the viscerocranial elements mainly serve to accommodate the water that was given a sufficient anterior-to-posterior impulse beforehand by powerful jaw separation. We hypothesize that this modified way of generating suction is primitive for salamanders, and that this behaviour could have played an important role in the evolution of terrestrial life in vertebrates by releasing mechanical constraints on the hyobranchial system.
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