Timothy E. Higham

The diversity of both the locomotor and feeding systems in fish is extensive, although little is known about the integrated evolution of the two systems. Virtually, all fish swim to ingest prey and all open their buccal cavity during prey capture, but the relationship between these two ubiquitous components of fish feeding strikes is unknown. We predicted that there should be a positive correlation between ram speed (RS) and maximum gape (MG) because the accuracy of a predatory strike goes down with an increase in RS and fish with larger mouths eat larger, more evasive prey. For 18 species of neotropical cichlids, we used phylogenetic‐independent contrasts to study the relationship between the predator closing speed (RS) and mouth size (MG) during prey capture. To provide a robust comparative framework, we augmented existing phylogenetic information available from the mitochondrial cytochrome b gene with sequences from the S7 nuclear ribosomal intron for these species. Then, we captured high‐speed (500 images per second), lateral view feeding sequences of each species by using a digital video camera and measured both RS and MG. Uncorrected species values of MG and RS were positively and significantly correlated. When accounting for any of the set of phylogenetic relationships recovered, the independent contrasts of RS and MG remained significantly, and positively, correlated. This tight evolutionary coupling highlights what is likely a common relationship between locomotor behaviour and feeding kinematics in many organisms.

SUMMARY: Suction feeding fish rapidly expand their oral cavity, resulting in a flow of water directed towards the mouth that is accompanied by a drop in pressure inside the buccal cavity. Pressure inside the mouth and fluid speed external to the mouth are understood to be mechanically linked but the relationship between them has never been empirically determined in any suction feeder. We present the first simultaneous measurements of fluid speed and buccal pressure during suction feeding in fishes. Digital particle image velocimetry (DPIV)and high-speed video were used to measure the maximum fluid speed in front of the mouth of four largemouth bass and three bluegill sunfish by positioning a vertical laser sheet on the mid-sagittal plane of the fish. Peak magnitude of pressure inside the buccal cavity was quantified using a transducer positioned within a catheter that opened into the dorsal wall of the buccal cavity. In both species the time of peak pressure preceded the time of peak fluid speed by as much as 42 ms, indicating a role for unsteady flow effects in shaping this relation. We parameterized an existing model of suction feeding to determine whether the relationship between peak pressures and fluid speeds that we observed could be predicted using just a few kinematic variables. The model predicted much higher fluid speeds than we measured at all values of peak pressure and gave a scaling exponent between them (0.51) that was higher than observed (0.36 for largemouth bass, 0.38 for bluegill). The scaling between peak buccal pressure and peak fluid speed at the mouth aperture differed in the two species, supporting the recent conclusion that species morphology affects this relation such that a general pattern may not hold.

SUMMARY: Suction feeding fish draw prey into the mouth using a flow field that they generate external to the head. In this paper we present a multidimensional perspective on suction feeding performance that we illustrate in a comparative analysis of suction feeding ability in two members of Centrarchidae, the largemouth bass (Micropterus salmoides) and bluegill sunfish(Lepomis macrochirus). We present the first direct measurements of maximum fluid speed capacity, and we use this to calculate local fluid acceleration and volumetric flow rate. We also calculated the ingested volume and a novel metric of strike accuracy. In addition, we quantified for each species the effects of gape magnitude, time to peak gape, and swimming speed on features of the ingested volume of water. Digital particle image velocimetry (DPIV) and high-speed video were used to measure the flow in front of the mouths of three fish from each species in conjunction with a vertical laser sheet positioned on the mid-sagittal plane of the fish. From this we quantified the maximum fluid speed (in the earthbound and fish's frame of reference), acceleration and ingested volume. Our method for determining strike accuracy involved quantifying the location of the prey relative to the center of the parcel of ingested water. Bluegill sunfish generated higher fluid speeds in the earthbound frame of reference, accelerated the fluid faster, and were more accurate than largemouth bass. However, largemouth bass ingested a larger volume of water and generated a higher volumetric flow rate than bluegill sunfish. In addition, because largemouth bass swam faster during prey capture, they generated higher fluid speeds in the fish's frame of reference. Thus, while bluegill can exert higher drag forces on stationary prey items, largemouth bass more quickly close the distance between themselves and prey. The ingested volume and volumetric flow rate significantly increased as gape increased for both species, while time to peak gape had little effect on the volume. However, peak gape distance did not affect the maximum fluid speed entering the mouth for either species. We suggest that species that generate high fluid speeds in the earthbound frame of reference will commonly exhibit small mouths and a high capacity to deliver force to buccal expansion,while species that ingest a large volume of water and generate high volumetric flow rates will have larger buccal cavities and cranial expansion linkage systems that favor displacement over force delivery.

Many natural animal movements involve accelerating from a standstill and then stopping. Obstacles in natural environments often limit the straight-line distance available for movement, and decreased braking ability theoretically can limit speed for short distances. Consequently, braking ability can be important for avoiding collisions with obstacles and exploiting resources effectively in complex environments. A presumed morphological correlate of improved braking performance in fish is increased pectoral fin area, because most fish protract these structures as they decelerate. However, the kinematics and modulation of velocity during starting and stopping are poorly understood for most species of fish as well as most species of animals. Thus, for bluegill sunfish Lepomis macrochirus with complete and partially ablated pectoral fins (35% original fin area), we analyzed high speed video recordings (200 images s^–1) of predatory attacks with a start and stop in a short, standardized distance (40 cm). We quantified body displacement, velocity, acceleration, deceleration and several fin angle variables during each feeding. Unexpectedly, several variables including maximum velocity and maximum deceleration (grand means 72 cm s^–1 and –512 cm s^–2, respectively) did not change significantly with reduced pectoral fin area. The average values of braking movements of the median and caudal fins did increase with decreased pectoral fin area but lacked statistically significant differences. The primary mechanism of attaining similar braking performance with decreased area of the pectoral fins was that they were protracted significantly more (mean difference=42°) and with a significantly faster average velocity of protraction. Thus, pectoral fin area appears unlikely to be the primary constraint on braking performance for this particular task.

It is well established that suction feeding fish use a variable amount of swimming (ram) during prey capture. However, the fluid mechanical effects of ram on suction feeding are not well established. In this study we quantified the effects of ram on the maximum fluid speed of the water entering the mouth during feeding as well as the spatial patterns of flow entering the mouth of suction-feeding bluegill sunfish Lepomis macrochirus. Using Digital Particle Image Velocimetry (DPIV) and high-speed video, we observed the flow in front of the mouth of three fish using a vertical laser sheet positioned on the mid-sagittal plane of the fish. From this we quantified the maximum fluid speed (measured at a distance in front of the mouth equal to one half of the maximum mouth diameter), the degree of focusing of water flow entering the mouth, and the shape of the ingested volume of water. Ram speed in 41 feeding sequences, measured at the time of maximum gape, ranged between 0 and 25 cm s^–1, and the ratio of ram speed to fluid speed ranged from 0.1% to 19.1%. In a regression ram speed did not significantly affect peak fluid speed, but with an increase in ram speed the degree of focusing of water entering the mouth increased significantly, and the shape of the ingested volume of water became more elongate and narrow. The implications of these findings are that (1) suction feeders that employ ram of between 0% and 20% of fluid speed sacrifice little in terms of the fluid speeds they generate and (2) ram speed enhances the total body closing speed of the predator.

SUMMARY Arboreal animals, especially lizards, often traverse three-dimensional networks of narrow perches with variable and steep inclines, but the effects of both incline and narrow surfaces on the locomotor movement and function of limbs are poorly understood. Thus, we quantified the three-dimensional hindlimb kinematics of a specialized arboreal lizard, Chamaeleo calyptratus, moving horizontally, and up and down a 30° incline on a narrow (2.4 cm) perch and a flat surface. We compared the flat-surface data of C. calyptratus with those of an anatomically generalized terrestrial lizard, Dipsosaurus dorsalis. Inclines had significant main effects for relatively few kinematic variables of C. calyptratus (11%) compared to D. dorsalis (73%). For C. calyptratus, the main effects of locomotor surface were nearly three times more widespread than those of incline. The foot of C. calyptratus was markedly anterior to the hip at footfall, primarily as a result of an unusually extended knee for a lizard. A large amount of knee flexion during early stance may be used by C. calyptratus to actively pull the body forward in a manner not found in D. dorsalis. Unexpectedly, the pelvic rotation of C. calyptratus greatly exceeded that of D. dorsalis and, unlike D. dorsalis, was not affected by incline. The more medial location of the foot of C. calyptratus on the narrow perch during stance was primarily a result of knee flexion rather than femur depression. Unlike previous qualitative descriptions of chameleons, our data for the hindlimb posture of C. calyptratus during stance indicate that the limb was not particularly erect.

SUMMARY Arboreal animals often move on surfaces with variable and steep inclines, but the changes in hindlimb muscle activity in response to incline are poorly understood. Thus, we studied the hindlimb muscle activity in the arboreal specialist, Chamaeleo calyptratus, moving up and down 45° inclines and on a horizontal surface. We quantified electromyograms (EMGs) from nine hindlimb muscles, and correlated EMGs with three-dimensional hindlimb kinematics. Kinematics changed little with incline, but the EMGs changed substantially. Most of the changes in EMGs were for amplitude rather than timing, and the EMGs of the hip and thigh muscles had more conspicuous changes with incline than those of the lower limb muscles. Unlike most other vertebrates, chameleons flexed the knee substantially during the first half of stance while the foot was anchored to the perch, and the amplitude of two large knee flexors increased when moving uphill compared to level and downhill. Thus, knee flexion in early stance probably contributes significantly to propulsion in C. calyptratus. During stance, the caudofemoralis EMGs of C. calyptratus correlated well with femur retraction, knee flexion and posterior femur rotation, and their amplitudes were higher on uphill and level surfaces than on the downhill surface. During the second half of stance, iliotibialis EMGs correlated well with knee extension, and their amplitude was highest on the uphill surface and lowest on the downhill surface. Many of the muscles in the hindlimb of C. calyptratus changed activity with incline in a manner similar to the propulsive limb muscles in mammals. Although muscle strain often increases when animals need more power to move uphill, the minimal changes in the hindlimb kinematics of C. calyptratus with incline imply little change in muscle strain.