The benefits of planar circular mouths on suction feeding performance

Suction feeding is the most common form of prey capture across aquatic feeding vertebrates and many adaptations that enhance efficiency and performance are expected. Many suction feeders have mechanisms that allow the mouth to form a planar and near-circular opening that is believed to have beneficial hydrodynamic effects. We explore the effects of the flattened and circular mouth opening through computational fluid dynamics simulations that allow comparisons with other mouth profiles. Compared to mouths with lateral notches, we find that the planar mouth opening results in higher flow rates into the mouth and a region of highest flow that is positioned at the centre of the mouth aperture. Planar mouths provide not only for better total fluid flow rates through the mouth but also through the centre of the mouth near where suction feeders position their prey. Circular mouths are shown to provide the quickest capture times for spherical and elliptical prey because they expose the prey item to a large region of high flow. Planar and circular mouths result in higher flow velocities with peak flow located at the centre of the mouth opening and they maximize the capacity of the suction feeders to exert hydrodynamic forces on the prey.     

The forces exerted by aquatic suction feeders on their prey

Successful prey capture by aquatic suction feeders depends on the ability of the predator to generate a flow of water external to the mouth that overcomes any movements and forces that the prey uses to resist the suction flow. Elucidating the nature and magnitude of these forces is a key to understanding what limits suction feeding performance. We identify three potential forces produced by the suction flow field: drag, acceleration reaction and the fluid pressure gradient. Using a mathematical model parametrized with empirical data from feeding bluegill, Lepomis macrochirus, we explore the relative magnitude of these forces under three encounter scenarios with a 5mm diameter, spherical prey: an immobile mid-water prey; a similar prey that executes an escape response; and a prey item that grips a substratum. Contrary to the almost exclusive emphasis on drag in the suction feeding literature, it made a minor contribution to the total forces in all three cases. In all three scenarios, the pressure gradient is the largest of the three forces. These results are important because previous researchers have emphasized drag and have not explicitly recognized a role for the pressure gradient force in suction feeding. The simulations suggest previously unrecognized mechanisms that suction feeders can use to enhance the forces that they exert, by increasing the steepness of the pressure gradient that the prey item is exposed to. This can be accomplished either by increasing the rate of increase in fluid velocity or by restricting the size of the mouth aperture, which creates a steeper spatial gradient in pressure.     

Relative importance of growth and behaviour to elasmobranch suction-feeding performance over early ontogeny.

Development of the ability to capture prey is crucial to predator survival. Trends in food-capture performance over early ontogeny were quantified for leopard sharks Triakis semifasciata and whitespotted bamboosharks Chiloscyllium plagiosum by measuring suction pressure and flow in front of the mouth during feeding. At any size, C. plagiosum produce greater subambient pressure and ingest more rounded water parcels. Maximum subambient pressure scaled with negative allometry in T. semifasciata and was accompanied by an increase in the time to reach maximum gape. Despite a similar trend in buccal expansion timing, maximum pressure in C. plagiosum scaled with isometry and was accompanied by an earlier onset of hyoid depression and a positive allometric increase in buccal reserve volume. Growth was the primary factor responsible for developmental trends in both species, with size-independent behavioural changes contributing little to overall performance variability. Ontogenetic dietary shifts are predicted for both species as a consequence of size-dependent changes in performance. Chiloscyllium plagiosum becomes anatomically and behaviourally canalized towards suction feeding, limiting the effective range of prey capture and possibly necessitating stalking. Triakis semifasciata, by contrast, retains the flexibility to employ both ram and suction and therefore captures more elusive prey with age.     

Use of computational fluid dynamics to study forces exerted on prey by aquatic suction feeders

Suction feeding is the most commonly used mechanism of prey capture among aquatic vertebrates. Most previous models of the fluid flow caused by suction feeders involve making several untested assumptions. In this paper, a Chimera overset grids approach is used to solve the governing equations of fluid dynamics in order to investigate the assumptions that prey do not interact with the flow and that the flow can be modelled as a one-dimensional flow. Results show that, for small prey, both neglecting the prey and considering prey interaction give similar calculated forces exerted on the prey. However, as the prey item increases in size toward the size of the gape, its effect on the flow becomes more pronounced. This in turn affects both the magnitude of the hydrodynamic forces imparted to the prey and the time when maximum force is delivered. Maximum force is delivered most quickly to intermediate sized prey, about one-third of mouth diameter, and most slowly to prey less than 7 per cent or greater than 67 per cent of mouth diameter. This suggests that the effect of prey size on the timing of suction forces may have substantial consequences for the feeding ecology of suction feeders that are known to prefer prey between 25 and 50 per cent of mouth diameter. Moreover, for a 15 cm fish with a 15 mm gape, assuming a radial one-dimensional flow field can result in underestimating the maximum force exerted on a 5 mm diameter spherical prey 1 gape distance from the mouth by up to 28.7 per cent.     

Anterior-to-posterior wave of buccal expansion in suction feeding fishes is critical for optimizing fluid flow velocity profile

In fishes that employ suction feeding, coordinating the timing of peak flow velocity with mouth opening is likely to be an important feature of prey capture success because this will allow the highest forces to be exerted on prey items when the jaws are fully extended and the flow field is at its largest. Although it has long been known that kinematics of buccal expansion in feeding fishes are characterized by an anterior-to-posterior wave of expansion, this pattern has not been incorporated in most previous computational models of suction feeding. As a consequence, these models have failed to correctly predict the timing of peak flow velocity, which according to the currently available empirical data should occur around the time of peak gape. In this study, we use a simple fluid dynamic model to demonstrate that the inclusion of an anterior-to-posterior wave of buccal expansion can correctly reproduce the empirically determined flow velocity profile, although only under very constrained conditions, whereas models that do not allow this wave of expansion inevitably predict peak velocity earlier in the strike, when the gape is less than half of its maximum. The conditions that are required to produce a realistic velocity profile are as follows: (i) a relatively long time lag between mouth opening and expansion of the more posterior parts of the mouth, (ii) a short anterior portion of the mouth relative to more posterior sections, and (iii) a pattern of movement that begins slowly and then rapidly accelerates. Greater maximum velocities were generated in simulations without the anterior-to-posterior wave of expansion, suggesting a trade-off between maximizing fluid speed and coordination of peak fluid speed with peak gape.     

Extremely fast prey capture in pipefish is powered by elastic recoil.

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(-1)), 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.     

Volumetric quantification of fluid flow reveals fish's use of hydrodynamic stealth to capture evasive prey

In aquatic ecosystems, predation on zooplankton by fish provides a major pathway for the transfer of energy to higher trophic levels. Copepods are an abundant zooplankton group that sense hydromechanical disturbances produced by approaching predators and respond with rapid escapes. Despite this capability, fish capture copepods with high success. Previous studies have focused on the predatory strike to elucidate details of this interaction. However, these raptorial strikes and resulting suction are only effective at short range. Thus, small fish must closely approach highly sensitive prey without triggering an escape in order for a strike to be successful. We use a new method, high-speed, infrared, tomographic particle image velocimetry, to investigate three-dimensional fluid patterns around predator and prey during approaches. Our results show that at least one planktivorous fish (Danio rerio) can control the bow wave in front of the head during the approach and consumption of prey (copepod). This alters hydrodynamic profiles at the location of the copepod such that it is below the threshold required to elicit an escape response. We find this behaviour to be mediated by the generation of suction within the buccopharyngeal cavity, where the velocity into the mouth roughly matches the forward speed of the fish. These results provide insight into how animals modulate aspects of fluid motion around their bodies to overcome escape responses and enhance prey capture.     

Aquatic suction feeding dynamics: insights from computational modelling

Aquatic suction feeding in vertebrates involves extremely unsteady flow, externally as well as internally of the expanding mouth cavity. Consequently, studying the hydrodynamics involved in this process is a challenging research area, where experimental studies and mathematical models gradually aid our understanding of how suction feeding works mechanically. Especially for flow patterns inside the mouth cavity, our current knowledge is almost entirely based on modelling studies. In the present paper, we critically discuss some of the assumptions and limitations of previous analytical models of suction feeding using computational fluid dynamics.     

Hydrodynamic modelling of aquatic suction performance and intra-oral pressures: limitations for comparative studies.

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.     

Rapid pivot feeding in pipefish: flow effects on prey and evaluation of simple dynamic modelling via computational fluid dynamics

Most theoretical models of unsteady aquatic movement in organisms assume that including steady-state drag force and added mass approximates the hydrodynamic force exerted on an organism''s body. However, animals often perform explosively quick movements where high accelerations are realized in a few milliseconds and are followed closely by rapid decelerations. For such highly unsteady movements, the accuracy of this modelling approach may be limited. This type of movement can be found during pivot feeding in pipefish that abruptly rotate their head and snout towards prey. We used computational fluid dynamics (CFD) to validate a simple analytical model of cranial rotation in pipefish. CFD simulations also allowed us to assess prey displacement by head rotation. CFD showed that the analytical model accurately calculates the forces exerted on the pipefish. Although the initial phase of acceleration changes the flow patterns during the subsequent deceleration phase, the accuracy of the analytical model was not reduced during this deceleration phase. Our analysis also showed that prey are left approximately stationary despite the quickly approaching pipefish snout. This suggests that pivot-feeding fish need little or no suction to compensate for the effects of the flow induced by cranial rotation.     

Jaw protrusion enhances forces exerted on prey by suction feeding fishes

The ability to protrude the jaws during prey capture is a hallmark of teleost fishes, widely recognized as one of the most significant innovations in their diverse and mechanically complex skull. An elaborated jaw protrusion mechanism has independently evolved multiple times in bony fishes, and is a conspicuous feature in several of their most spectacular radiations, ultimately being found in about half of the approximately 30 000 living species. Variation in jaw protrusion distance and speed is thought to have facilitated the remarkable trophic diversity found across fish groups, although the mechanical consequences of jaw protrusion for aquatic feeding performance remain unclear. Using a hydrodynamic approach, we show that rapid protrusion of the jaws towards the prey, coupled with the spatial pattern of the flow in front of the mouth, accelerates the water around the prey. Jaw protrusion provides an independent source of acceleration from that induced by the unsteady flow at the mouth aperture, increasing by up to 35% the total force exerted on attached, escaping and free-floating passive prey. Despite initiating the strike further away, fishes can increase peak force on their prey by protruding their jaws towards it, compared with a ‘non-protruding’ state, where the distance to prey remains constant throughout the strike. The force requirements for capturing aquatic prey might have served as a selective factor for the evolution of jaw protrusion in modern fishes.     

A minimal model of predator–swarm interactions

We propose a minimal model of predator–swarm interactions which captures many of the essential dynamics observed in nature. Different outcomes are observed depending on the predator strength. For a ‘weak’ predator, the swarm is able to escape the predator completely. As the strength is increased, the predator is able to catch up with the swarm as a whole, but the individual prey is able to escape by ‘confusing’ the predator: the prey forms a ring with the predator at the centre. For higher predator strength, complex chasing dynamics are observed which can become chaotic. For even higher strength, the predator is able to successfully capture the prey. Our model is simple enough to be amenable to a full mathematical analysis, which is used to predict the shape of the swarm as well as the resulting predator–prey dynamics as a function of model parameters. We show that, as the predator strength is increased, there is a transition (owing to a Hopf bifurcation) from confusion state to chasing dynamics, and we compute the threshold analytically. Our analysis indicates that the swarming behaviour is not helpful in avoiding the predator, suggesting that there are other reasons why the species may swarm. The complex shape of the swarm in our model during the chasing dynamics is similar to the shape of a flock of sheep avoiding a shepherd.     

Prey should hide more randomly when a predator attacks more persistently

When being searched for and then (if found) pursued by a predator, a prey animal has a choice between choosing very randomly among hiding locations so as to be hard to find or alternatively choosing a location from which it is more likely to successfully flee if found. That is, the prey can choose to be hard to find or hard to catch, if found. In our model, capture of prey requires both finding it and successfully pursuing it. We model this dilemma as a zero-sum repeated game between predator and prey, with the eventual capture probability as the pay-off to the predator. We find that the more random hiding strategy is better when the chances of repeated pursuit, which are known to be related to area topography, are high. Our results extend earlier results of Gal and Casas, where there was at most only a single pursuit. In that model, hiding randomly was preferred by the prey when the predator has only a few looks. Thus, our new multistage model shows that the effect of more potential looks is opposite. Our results can be viewed as a generalization of search games to the repeated game context and are in accordance with observed escape behaviour of different animals.     

The smaller your mouth, the longer your snout: predicting the snout length of Syngnathus acus, Centriscus scutatus and other pipette feeders

Like most ray-finned fishes (Actinopterygii), pipefishes (Syngnathoidei) feed by suction. Most pipefishes reach their prey by a rapid dorso-rotation of the head. In the present study, we analysed the feeding kinematics of the razor fish, Centriscus scutatus, and of the greater pipefish, Syngnathus acus in detail. We found capture times of as little as 4-6ms for C. scutatus and 6-8ms for S. acus. We then hypothesized that the long snout of pipefishes is optimal for such fast feeding. To test this, we implemented in a mathematical model the following considerations. To reach the prey as fast as possible, a low moment of inertia increases the head's angular speed, whereas a long snout decreases the angle over which the head must be turned. The model accurately predicted the snout lengths of a number of pipefishes. We found that the optimal snout length, with which a prey will be reached fastest, is inversely related to its cross-section. In spite of the small cross-section, the development of a long snout can be an evolutionary advantage because this reduces the time to approach the prey.     

具有时滞和脉冲捕获、污染物脉冲输入的捕食-食饵-环境模型

本文建立了污染环境中,捕食者具有阶段结构、食饵具有脉冲收获、污染物具有脉冲输入的时滞捕食-食饵-环境模型,利用离散动力系统的频闪映射和比较原理,得到了捕食者灭绝周期解的全局吸引性和系统持久性的充分条件．通过数值仿真,研究了食饵的捕获、污染物的脉冲输入和脉冲作用周期对捕食者灭绝和持续生存的影响,同时也验证了理论结果．对生物资源的开发、种群数量的收获及环境的控制提供了宝贵的理论依据．     

Hydrodynamic constraints on prey-capture performance in forward-striking snakes

Some specialized aquatic snakes such as Natrix tessellata strike at fish by rapidly accelerating their head towards the prey with their mouth opened widely. This strategy is believed to be suboptimal as relatively high drag forces act on the open jaws and, therefore, probably limit strike speed. Moreover, the bow wave in front of the snake''s jaws could push prey away from the mouth, thus potentially explaining the relatively low capture success observed in these animals (<20%). Here, we used laser-scan based computational fluid dynamics to test these potential constraints on prey-capture performance for N. tessellata. Our simulations showed that drag force indeed increases drastically for striking at a high gape angle. However, we estimated the overall cost in slowing down strike speed to be less pronounced due to the instationary dynamics of the system. In contrast to the expectations, forward displacement of prey was relatively limited (<13% of head length), and forceful collisions between prey and the leading edge of the jaw regularly occurred. However, our models showed that precise aiming by the snake was needed to reduce the chance of deviating the prey to a path bypassing the mouth. Our study also indicated several hydrodynamic advantages for snakes to strike at relatively large prey.     

Modelled three-dimensional suction accuracy predicts prey capture success in three species of centrarchid fishes

Prey capture is critical for survival, and differences in correctly positioning and timing a strike (accuracy) are likely related to variation in capture success. However, an ability to quantify accuracy under natural conditions, particularly for fishes, is lacking. We developed a predictive model of suction hydrodynamics and applied it to natural behaviours using three-dimensional kinematics of three centrarchid fishes capturing evasive and non-evasive prey. A spheroid ingested volume of water (IVW) with dimensions predicted by peak gape and ram speed was verified with known hydrodynamics for two species. Differences in capture success occurred primarily with evasive prey (64–96% success). Micropterus salmoides had the greatest ram and gape when capturing evasive prey, resulting in the largest and most elongate IVW. Accuracy predicted capture success, although other factors may also be important. The lower accuracy previously observed in M. salmoides was not replicated, but this is likely due to more natural conditions in our study. Additionally, we discuss the role of modulation and integrated behaviours in shaping the IVW and determining accuracy. With our model, accuracy is a more accessible performance measure for suction-feeding fishes, which can be used to explore macroevolutionary patterns of prey capture evolution.     

Numerical Modelling of Flow Boiling Heat Transfer in Horizontal Metal‐Foam Tubes

The flow boiling heat transfer performance in horizontal metal‐foam tubes is numerically investigated based on the flow pattern map retrieved from experimental investigations. The flow pattern and velocity profile are generally governed by vapour quality and mass flow rate of the fluid. The porous media non‐equilibrium heat transfer model is employed for modelling both vapour and liquid phase zones. The modelling predictions have been compared with experimental results. The effects of metal‐foam morphological parameters, heat flux and mass flux on heat transfer have been examined. The numerical predictions show that the overall heat transfer coefficient of the metal‐foam filled tube increases with the relative density (1‐porosity), pore density (ppi), mass and heat flux.     

Predator confusion is sufficient to evolve swarming behaviour

Swarming behaviours in animals have been extensively studied owing to their implications for the evolution of cooperation, social cognition and predator–prey dynamics. An important goal of these studies is discerning which evolutionary pressures favour the formation of swarms. One hypothesis is that swarms arise because the presence of multiple moving prey in swarms causes confusion for attacking predators, but it remains unclear how important this selective force is. Using an evolutionary model of a predator–prey system, we show that predator confusion provides a sufficient selection pressure to evolve swarming behaviour in prey. Furthermore, we demonstrate that the evolutionary effect of predator confusion on prey could in turn exert pressure on the structure of the predator''s visual field, favouring the frontally oriented, high-resolution visual systems commonly observed in predators that feed on swarming animals. Finally, we provide evidence that when prey evolve swarming in response to predator confusion, there is a change in the shape of the functional response curve describing the predator''s consumption rate as prey density increases. Thus, we show that a relatively simple perceptual constraint—predator confusion—could have pervasive evolutionary effects on prey behaviour, predator sensory mechanisms and the ecological interactions between predators and prey.     

The effect of external flow on the feeding currents of sessile microorganisms

Microscopic sessile suspension feeders live attached to surfaces and, by consuming bacteria-sized prey and by being consumed, they form an important part of aquatic ecosystems. Their environmental impact is mediated by their feeding rate, which depends on a self-generated feeding current. The feeding rate has been hypothesized to be limited by recirculating eddies that cause the organisms to feed from water that is depleted of food particles. However, those results considered organisms in still water, while ambient flow is often present in their natural habitats. We show, using a point-force model, that even very slow ambient flow, with speed several orders of magnitude less than that of the self-generated feeding current, is sufficient to disrupt the eddies around perpendicular suspension feeders, providing a constant supply of food-rich water. However, the feeding rate decreases in external flow at a range of non-perpendicular orientations due to the formation of recirculation structures not seen in still water. We quantify the feeding flow and observe such recirculation experimentally for the suspension feeder Vorticella convallaria in external flows typical of streams and rivers.