A KINEMATIC STUDY OF SUCTION FEEDING AND ASSOCIATED BEHAVIOR IN THE LONG-FINNED PILOT WHALE, GLOBICEPHALA MELAS (TRAILL)

Analysis of videotaped feeding sequences provides novel documentation of suction feeding in captive juvenile long-finned pilot whales ( Globicephala melas ). Swimming and stationary whales were videotaped while feeding at the surface, mid-water, and bottom. The ingestion sequence includes a preparatory phase with partial gape followed by jaw opening and rapid hyoid depression to suck in prey at a mean distance of 14 cm (duration 90 msec), although prey were taken from much greater distances. Depression and retraction of the large, piston-like tongue generate negative intraoral pressures for prey capture and ingestion. Food was normally ingested without grasping by teeth yet was manipulated with lingual, hyoid, and mandibular movement for realignment; suction was then used to transport prey into the oropharynx. Whales frequently rolled or inverted before taking prey, presumably to avoid grasping and repositioning. Prey were sucked off the bottom or sides of the pool without direct contact; lateral suction was used to ingest items from the sides of the mouth.     

The roles of visual and proprioceptive information during motor program choice in frogs

Previous studies have shown that leopard frogs, Rana pipiens, use tongue prehension to capture small prey and jaw prehension to capture large prey. After hypoglossal nerve transection, the frogs fail to open their mouths when attempting to feed on small prey, but open their mouths and capture large prey. Here, we investigate how visual information about the prey and proprioceptive information from the tongue interact to influence the motor program choice. Using pieces of earthworm of various sizes, we found that Rana exhibits two different behavior patterns based on prey size. The frogs captured the 1.5-cm prey using tongue prehension, whereas 2.0-cm and larger prey were captured using jaw prehension. After hypoglossal transection, the frogs never opened their mouths when they tried to feed on 1.5-cm prey. When feeding on 3.0-cm and larger prey after transection, they always opened their mouths and captured the prey using jaw prehension. When offered 2.0-cm prey, they alternated randomly between opening and not opening the mouth. Therefore, deafferentation changed the pattern of motor program choice at the behavioral border. This implies that afferents from the tongue interact with visual input to influence motor program choice.     

Prey capture behavior and kinematics of the Atlantic cownose ray, Rhinoptera bonasus

The structurally reinforced jaws of the cownose ray, Rhinoptera bonasus testify to this species' durophagous diet of mollusks, but seem ill-suited to the behaviors necessary for excavating such prey. This study explores this discordance by investigating the prey excavation and capture kinematics of R. bonasus. Based on the basal suction feeding mechanism in this group of fishes, we hypothesized a hydraulic method of excavation. As expected, prey capture kinematics of R. bonasus show marked differences relative to other elasmobranchs, relating to prey excavation and use of the cephalic lobes (modified anterior pectoral fin extensions unique to derived myliobatiform rays). Prey are excavated by repeated opening and closing of the jaws to fluidize surrounding sand. The food item is then enclosed laterally by the depressed cephalic lobes, which transport it toward the mouth for ingestion by inertial suction. Unlike in most sharks, upper jaw protrusion and mandibular depression are simultaneous. During food capture, the ray's spiracle, mouth, and gill slit movements are timed such that water enters only the mouth (e.g., the spiracle closes prior to prey capture and reopens immediately following). Indigestible parts are then hydraulically winnowed from edible prey portions, by mouth movements similar to those used in excavation, and ejected through the mouth. The unique sensory/manipulatory capabilities of the cephalic lobes, as well as the cownose ray's hydraulic excavation/winnowing behaviors and suction feeding, make this species an effective benthic predator, despite its epibenthic lifestyle.     

Prey location,biomechanical constraints,and motor program choice during prey capture in the tomato frog, <Emphasis Type="Italic">Dyscophus guineti</Emphasis>

This study investigated how visual information about prey location and biomechanical constraints of the feeding apparatus influence the feeding behavior of the tomato frog, Dyscophus guineti. When feeding on prey at small azimuths (less than ± 40°), frogs aimed their heads toward the prey but did not aim their tongues relative to their heads. Frogs projected their tongues rapidly by transferring momentum from the lower jaw to the tongue. Storage and recovery of elastic energy by the mouth opening muscles amplified the velocities of mouth opening and tongue projection. This behavior can only occur when the lower jaw and tongue are aligned (i.e., within the range of motion of the neck). When feeding on prey at large azimuths (greater than ± 40°), frogs aimed both the head and tongue toward the prey and used a muscular hydrostatic mechanism to project the tongue. Hydrostatic elongation allows for frogs to capture prey at greater azimuthal locations. Because the tongue moves independently of the lower jaw, frogs can no longer take advantage of momentum transfer to amplify the speed of tongue projection. To feed on prey at different azimuthal locations, tomato frogs switch between alternative strategies to circumvent these biomechanical constraints.     

Piscivorous cyprinid fish modulates suction feeding kinematics to capture elusive prey

Previous studies have shown that evasive prey generally elicit a different kinematical pattern of prey capture from suction feeding fish compared to non-evasive types of prey. However, no evidence exists that predatory fish can modulate their prey capture kinematics in response to whether or not an elusive prey performs an escape response. Here, we analyse prey capture kinematics of a specialist piscivore (asp, Aspius aspius) during feeding on untethered, live goldfish, which regularly displayed escape attempts when attacked by the asp. Significant modulation occurred in function of the escape attempts of prey: mouth opening was prolonged and increased in magnitude, and one individual also showed an increased hyoid depression when feeding on prey trying to escape. As the orientation of the prey with respect to the predator prior to the start of mouth opening was related to the probability of observing an escape attempt, asp could theoretically perform this type of modulation by a priori choosing a pre-programmed motor pattern. However, since contact between the prey and the asp's mouth appeared to be a factor improving the timing of mouth closing, this fine-tuning of prey capture kinematics is more likely to be caused by reflexive neural feedback control.     

Tongue evolution in the lungless salamanders, family plethodontidae. I. Introduction, theory and a general model of dynamics.

Plethodontid salamanders capture prey by projecting the tongue from the mouth. An analysis of theoretical mechanics of the hyobranchial skeleton is used to formulate a working hypothesis of tongue movements. Predictions that the skeletal elements of the tongue are included in the projectile and that the hyobranchial skeleton is folded during projection are central to the analysis. When decapitated in a particular way, salamanders project the tongue, and it is not retracted. When these heads are fixed and sectioned, examination confirms the predications. In turn, these observations are used to refine the working hypothesis and to generate a general model of tongue dynamics for plethodontids. Muscles performing the major roles of projection (subarcualis rectus I) and retraction (rectus cervicis profundus) are identified. The skeleton is folded passively along a morphological track having the form of a tractrix. Predictions concerning the shape of the track and the exact configuration of the folded skeleton are confirmed by study of sectioned material. The skeleton unfolds along the track during retraction and is spread into the resting state. The model developed herein will be used as a basis for predictions concerning selection patterns in the family and for analytical purposes in comparative and evolutionary studies.     

Modulation of prey capture kinematics and the role of lingual sensory feedback in the lizard Pogona vitticeps

Most organisms feed on a variety of prey that may differ dramatically in their physical and behavioural characteristics (e.g. mobility, mass, texture, etc.). Thus the ability to modulate prey capture behaviour in accordance with the characteristics of the food appears crucial. In animals that use rapid tongue movements to capture prey (frogs and chameleons), the coordination of jaws and tongue is based on visual cues gathered prior to the prey capture event. However, most iguanian lizards have much slower tongue-based prey capture systems suggesting that sensory feedback from the tongue may play an important role in coordinating jaw and tongue movements. We investigated the modulation of prey capture kinematics in the agamid lizard Pogona vitticeps when feeding on a range of food items differing in their physical characteristics. As the lizard is a dietary generalist, we expected it to be able to modulate its prey capture kinematics as a function of the (mechanical) demands imposed by the prey. Additionally, we investigated the role of lingual sensory feedback by transecting the trigeminal sensory afferents. Our findings demonstrated that P. vitticeps modulates its prey capture kinematics according to specific prey properties (e.g. size). In addition, transection of the trigeminal sensory nerves had a strong effect on prey capture kinematics. However, significant prey type effects and prey type by transection effects suggest that other sources of sensory information are also used to modulate the prey capture kinematics in P. vitticeps.     

THE EVOLUTION OF TETRAPOD FEEDING BEHAVIOR: KINEMATIC HOMOLOGIES IN PREY TRANSPORT

One of the major features of the aquatic-to-terrestrial transition in vertebrate evolution was the change in the mechanism used to transport prey from the jaws to the throat. Primarily, vertebrates use hydraulic transport, but the transition to terrestrial life was accompanied by modifications of the hyobranchial apparatus that permit tongue-based transport. Despite an extensive data base on amniote feeding systems and mechanisms of intraoral prey transport, few data are available on the mechanism of prey transport in anamniote tetrapods. Transport cycles of four Ambystoma tigrinum (Amphibia) feeding on worms and crickets were filmed at 150 flames per second to produce quantitative profiles of the intraoral transport cycles for the two prey types. During the transport cycle the head and body remain stationary relative to the background: transport in Ambystoma tigrinum thus does not involve inertial movements of the head or body. Prey type had little effect on the kinematics of prey transport. The process of prey transport may be divided into four phases: preparatory, fast opening, closing, and recovery. The preparatory phase itself is divided into two parts: an extended segment that may include slight slow opening and a static phase prior to mouth opening where no change in gape occurs. The kinematic profile of transport in terrestrial salamanders is extremely similar to that used by fishes during hydraulic (aquatic) prey transport. We hypothesize that the distinct recovery and preparatory phases in the transport cycle of anamniote tetrapods are together homologous to the slow opening phases of the amniote cycle, and that during the evolution of terrestrial prey processing systems the primitive extended preparatory phase has become greatly compressed and incorporated into the amniote gape cycle.     

Movement of the hyoid in frogs during feeding.

Feeding, breathing, and vocalization sequences of Bufo marinus were recorded by cineradiography. Results of film analysis indicate that the hyoid moves during all three behaviors. Movement of the hyoid is critical in tongue protrusion of frogs, and a biomechanical model of this action is presented. The hyoid appears to represent a compromise morphological system for three functions, rather than an optimal system for any one. This may explain, in part, the retention of a relatively inefficient breathing mechanism in frogs.     

Functional morphology of the feeding mechanism in aquatic ambystomatid salamanders

This study addresses four questions in vertebrate functional morphology through a study of aquatic prey capture in ambystomatid salamanders: (1) How does the feeding mechanism of aquatic salamanders function as a biomechanical system? (2) How similar are the biomechanics of suction feeding in aquatic salamanders and ray-finned fishes? (3) What quantitative relationship does information extracted from electromyograms of striated muscles bear to kinematic patterns and animal performance? and (4) What are the major structural and functional patterns in the evolution of the lower vertebrate skull? During prey capture, larval ambystomatid salamanders display a kinematic pattern similar to that of other lower vertebrates, with peak gape occurring prior to both peak hyoid depression and peak cranial elevation. The depressor mandibulae, rectus cervicis, epaxialis, hypaxialis, and branchiohyoideus muscles are all active for 40–60 msec during the strike and overlap considerably in activity. The two divisions of the adductor mandibulae are active in a continuous burst for 110–130 msec, and the intermandibularis posterior and coracomandibularis are active in a double burst pattern. The antagonistic depressor mandibulae and adductor mandibulae internus become active within 0.2 msec of each other, but the two muscles show very different spike and amplitude patterns during their respective activity periods. Coefficients of variation for kinematic and most electromyographic recordings reach a minimum within a 10 msec time period, just after the mouth starts to open. Pressure within the buccal cavity during the strike reaches a minimum of ?25 mmHg, and minimum pressure occurs synchronously with maximum gill bar adduction. The gill bars (bearing gill rakers that interlock with rakers of adjacent arches) clearly function as a resistance within the oral cavity and restrict posterior water influx during mouth opening, creating a unidirectional flow during feeding. Durations of electromyographic activity alone are poor predictors of kinematic patterns. Analyses of spike amplitude explain an additional fraction of the variance in jaw kinematics, whereas the product of spike number and amplitude is the best statistical predictor of kinematic response variables. Larval ambystomatid salamanders retain the two primitive biomechanical systems for opening and closing the mouth present in nontetrapod vertebrates: elevation of the head by the epaxialis and depression of the mandible by the hyoid apparatus.     

The functional morphology of lingual prey capture in a scincid lizard,Tiliqua scincoides (Reptilia: Squamata)

We investigated the functional morphology of lingual prey capture in the blue‐tongued skink, Tiliqua scincoides, a lingual‐feeding lizard nested deep within the family Scincidae, which is presumed to be dominated by jaw‐feeding. We used kinematic analysis of high‐speed video to characterize jaw and tongue movements during prey capture. Phylogenetically informed principal components analysis of tongue morphology showed that, compared to jaw‐feeding scincids and lacertids, T. scincoides and another tongue‐feeding scincid, Corucia zebrata, are distinct in ways suggesting an enhanced ability for hydrostatic shape change. Lingual feeding kinematics show substantial quantitative and qualitative variation among T. scincoides individuals. High‐speed video analysis showed that T. scincoides uses significant hydrostatic elongation and deformation during protrusion, tongue‐prey contact, and retraction. A key feature of lingual prey capture in T. scincoides is extensive hydrostatic deformation to increase the area of tongue‐prey contact, presumably to maximize wet adhesion of the prey item. Adhesion is mechanically reinforced during tongue retraction through formation of a distinctive “saddle” in the foretongue that supports the prey item, reducing the risk of prey loss during retraction.     

The role of hypoglossal sensory feedback during feeding in the marine toad, Bufo marinus.

Behavioral observations demonstrate that bilateral deafferentation of the hypoglossal nerves in the marine toad (Bufo marinus) prevents mouth opening during feeding. In the present study, we used high-speed videography, electromyography (EMG), deafferentation, muscle stimulation, and extracellular recordings from the trigeminal nerve to investigate the mechanism by which sensory feedback from the tongue controls the jaw muscles of toads. Our results show that sensory feedback from the tongue enters the brain through the hypoglossal nerve during normal feeding. This feedback appears to inhibit both tonic and phasic activity of the jaw levators. Hypoglossal feedback apparently functions to coordinate tongue protraction and mouth opening during feeding. Among anurans, the primitive condition is the absence of a highly protrusible tongue and the absence of a hypoglossal sensory feedback system. The hypoglossal feedback system evolved in parallel with the acquisition of a highly protrusible tongue in toads and their relatives.     

The effects of intraspecific competition on the prey capture behavior and kinematics of the bluegill sunfish, Lepomis macrochirus

Competition has broad effects on fish and specifically the effects of competition on the prey capture kinematics and behavior are important for the assessment of future prey capture studies in bony fishes. Prey capture kinematics and behavior in bony fishes have been shown to be affected by temperature and satiation. The densities at which bony fish are kept have also been shown to affect their growth, behavior, prey selection, feeding and physiology. We investigated how density induced intraspecific competition for food affects the prey capture kinematics of juvenile bluegill sunfish, Lepomis macrochirus. High speed video was utilized to film five bold individuals feeding at three different densities representing different levels of intraspecific competition. We hypothesized that: (1) the feeding kinematics will be faster at higher levels of competition compared to lower levels of competition, and (2) bluegill should shift from more suction-based feeding towards more ram-based feeding with increasing levels of competition in order to outcompete conspecifics for a prey item. We found that, with increased intraspecific competition, prey capture became faster, involving more rapid jaw opening and therefore greater inertial suction, shorter mouth closing times, and shorter gape cycles. Furthermore, the attack velocity of the fish increased with increasing competition, however a shift towards primarily ram based feeding was not confirmed. Our study demonstrates that prey capture kinematics are affected by the presence of conspecifics and future studies need to consider the effects of competition on prey capture kinematics.     

Kinematics of the jaw and hyolingual apparatus during feeding in Caiman crocodilus

Movements of the neck, jaws, and hyolingual apparatus during inertial feeding in Caiman crocodilus were studied by cineradiography. Analysis reveals two kinds of cycles: inertial bites (reposition, kill/crush, and transport) and swallowing cycles. They differ in their gape profile and in displacement of the neck, cranium, and hyolingual apparatus. Inertial bites are initiated by an elevation of the neck and cranium; the head is then retracted backward, the prey simultaneously being lifted by the hyolingual apparatus. Next the lower jaw is depressed, and the prey is rapidly pushed further upward by the hyolingual apparatus. Thereafter fast mouth-closure occurs with the neck and cranium being abruptly depressed, the lower jaw elevated, and the hyolingual apparatus rapidly retracted ventrally. Depression of the neck and cranium thrusts the head forward and impacts the backward moving prey more posteriorly in the oral cavity. Swallowing cycles initially involve movement of the hyoid in front of the prey followed by rapid posteroventrad retraction of the hyoid, forcing the prey into the esophagus during opening and closing of the mouth. After mouth-closure, the hyoid apparatus is again protracted. Jaws, neck, tongue, and hyoid apparatus play an active role during intertial feeding sequences. At the beginning of a feeding sequence, the hyolingual apparatus mainly moves dorsoventrally, whereas toward the end of a sequence anteroposterior displacements of the hyoid are prominent. © 1992 Wiley-Liss, Inc.     

Predicting patterns of prey use from morphology of fishes

Synopsis Ecomorphological analyses that search for patterns of association between morphological and prey-use data sets will have a greater chance of understanding the causal relationships between form and diet if the morphological variables used have known consequences for feeding performance. We explore the utility of fish body size, mouth gape and jaw-lever mechanics in predicting patterns of prey use in two very different communities of fishes, Caribbean coral reef fishes, and species of the Centrarchidae that live in Lake Opinicon, Ontario. In spite of major differences in the spectrum of potential prey available, the centrarchids of Lake Opinicon show dietary transitions during ontogeny that are very similar to those seen among and within species of Caribbean groupers (Serranidae). The transition from small zooplankton to intermediate sized invertebrates and ultimately to fishes appears to be very general in ram-suction feeding fishes and is probably driven largely by the constraints of mouth size on prey capture ability. The jaw-lever systems for mouth opening and closing represent direct trade-offs for speed and force of jaw movement. The ratio of in-lever to out-lever in the opening system changes during ontogeny in bluegill, indicating that the mechanics and kinematics of jaw movement may change as well. Among 34 species of Caribbean reef fishes, biting species had jaw-closing ratios that favored force translation, while species that employ rapid-strike ram-suction had closing ratios that enhanced speed of closing and mouth opening ratios that favored a more rapid expansion of the mouth during the strike. We suggest that when prey are categorized into functional groups, reflecting the specific performance features that are important in capturing and handling them, and the differences among habitats in the available prey resource are taken into account, general patterns can be found in morphology-diet relations that cross phylogenetic boundaries.     

Cold-blooded snipers: thermal independence of ballistic tongue projection in the salamander Hydromantes platycephalus

Plethodontid salamanders of the genus Hydromantes capture prey using the most extreme tongue projection among salamanders, and can shoot the tongue a distance of 80% of body length in less than 20?msec. The tongue skeleton is projected from the body via an elastic-recoil mechanism that decouples muscle contraction from tongue projection, amplifying muscle power tenfold. We tested the hypothesis that the elastic-recoil mechanism also endows tongue projection with low thermal dependence by examining the kinematics and dynamics of tongue projection in Hydromantes platycephalus over a range of body temperatures (2-24°C). We found that H. platycephalus maintained tongue-projection performance over the tested temperature range and that tongue projection showed thermal independence (Q(10) values of 0.94-1.04) of all performance parameters including projection distance, average velocity, and peak instantaneous values of velocity, acceleration, and power. Nonelastic, muscle-powered tongue retraction, in contrast, responded to temperature changes significantly differently than elastic tongue projection; performance parameters of retraction displayed thermal dependence typical of muscle-powered movement (Q(10) values of 1.63-4.97). These results reveal that the elastic-recoil mechanism liberates tongue projection from the effects of temperature on muscle contractile rates. We suggest that relative thermal independence is a general characteristic of elastic-recoil mechanisms and may promote the evolution of these mechanisms in ectothermic animals.     

Morphology and function of the feeding apparatus of the lungfish, Lepidosiren paradoxa (Dipnoi)

The feeding mechanism of the South American lungfish, Lepidosiren paradoxa retains many primitive teleostome characteristics. In particular, the process of initial prey capture shares four salient functional features with other primitive vertebrates: 1) prey capture by suction feeding, 2) cranial elevation at the cranio-vertebral joint during the mouth opening phase of the strike, 3) the hyoid apparatus plays a major role in mediating expansion of the oral cavity and is one biomechanical pathway involved in depressing the mandible, and 4) peak hyoid excursion occurs after maximum gape is achieved. Lepidosiren also possesses four key morphological and functional specializations of the feeding mechanism: 1) tooth plates, 2) an enlarged cranial rib serving as a site for the origin of muscles depressing the hyoid apparatus, 3) a depressor mandibulae muscle, apparently not homologous to that of amphibians, and 4) a complex sequence of manipulation and chewing of prey in the oral cavity prior to swallowing. The depressor mandibulae is always active during mouth opening, in contrast to some previous suggestions. Chewing cycles include alternating adduction and transport phases. Between each adduction, food may be transported in or out of the buccal cavity to position it between the tooth plates. The depressor mandibulae muscle is active in a double-burst pattern during chewing, with the larger second burst serving to open the mouth during prey transport. Swallowing is characterized by prolonged activity in the hyoid constrictor musculature and the geniothoracicus. Lepidosiren uses hydraulic transport achieved by movements of the hyoid apparatus to position prey within the oral cavity. This function is analogous to that of the tongue in many tetrapods.     

Lamina-associated polypeptide 2 (LAP2) expression in fish and amphibians

Somatic and germinal cells of 15 fish and 33 amphibian species were examined by SDS-PAGE followed by immunoblotting to determine the expression of LAP2 (lamina-associated polypeptide 2). LAP2 expression in frogs, salamanders and fish does not vary with the mode of reproduction. In fish and frog cells, a rim-like LAP2 positive region was detected around the nucleus by indirect immunofluorescence microscopy. The cell distribution and expression patterns of LAP2 in fish, frogs and salamanders are comparable with those found in Xenopus and zebrafish. The mammalian somatic cell pattern, which may also occur in gymnophione amphibians, includes LAP2alpha, beta and gamma as major isoforms, whereas LAP2alpha does not occur in cells of fish, frogs and salamanders. In fish, LAP2gamma is the major isoform of somatic cells, suggesting that LAP2gamma may be ancestral. However, in the rainbow trout, as in frogs and salamanders, LAP2beta was the major somatic isoform. Fish and frog sperm only express low molecular weight polypeptides. In contrast, fish and frog oocytes express an oocyte-specific LAP2 isoform of high molecular weight. In the toad Bufo marinus this isoform becomes upregulated in pre-vitellogenic oocytes of 150-200 microm in diameter. The absence of LAP2alpha and the differential expression of LAP2 isoforms in somatic and germ cells, as found in fish and frogs, may be ancestral vertebrate characters. In spite of differences in developmental time, the LAP2 isoforms of somatic cells are upregulated during gastrulation, suggesting that LAP2 may be implicated in the early development of fish and frog.     

Testing amniote models of prey transport kinematics: a quantitative analysis of mouth opening patterns in lizards

Two models have been proposed to describe the prey transport kinematics of terrestrial vertebrates (Bramble and Wake, 1985; Reilly and Lauder, 1990). The critical difference between the models is the presence or absence of a slow open-II phase (SO-II) in the gape profile during mouth opening. Each of these models has been applied to lizards, however to date, lizard feeding kinematics have not been adequately quantified to assess the utility of these models for this clade. Neither model has been sufficiently tested due to the lack of a methodology to assess the specific differences between the models. We describe a method that uses explicit mathematical criteria to define the kinematic phases in tetrapod feeding. This "slope analysis& is used to precisely quantify and compare the transport kinematics of seven lizard species. Lizard transport kinematics were highly variable both within and across taxa. However, several common gape cycle patterns were identified. The predominant patterns were slow-fast opening (37.3%), fast opening only (22.9%) and slow opening only (21.2%). The most common pattern explicitly fits the prediction of the Reilly and Lauder model while the other two are similar to patterns observed in salamanders. Thus, lizards possess both the slow opening-fast opening pattern predicted for amniotes and the more primitive, simple opening pattern characteristic of more basal tetrapods. Plateau phases were found in only 12.8% of the profiles and only a fourth of these (3.4% of the total) explicitly fit the Bramble and Wake model (slow opening, plateau, fast opening) and two species never exhibited plateaus in their gape cycles. Thus, it is clear that the Bramble and Wake model is not supported as a generalized model for lizards or generalized tetrapods.     

Kinetics of tongue projection in Ambystoma tigrinum: Quantitative kinematics,muscle function,and evolutionary hypotheses

The projectile tongue of caudate amphibians has been studied from many perspectives, yet a quantitative kinetic model of tongue function has not yet been presented for generalized (nonplethodontid) terrestrial salamanders. The purposes of this paper are to describe quantitatively the kinnematics of the feeding mechanism and to present a kinetic model for the function of the tongue in the ambystomatid salamander Ambystoma tigrinum. Six kinematic variables were quantified from high-speed films of adult A. tigrinum feeding on land and in the water. Tongue protrusion reaches its maximum during peak gape, while peak tongue height is reached earlier, 15 ms after the mouth starts to open. Tongue kinematics change considerably during feeding in the water, and the tongue is not protruded past the plane of the gape. Electrical stimulation of the major tongue muscles showed that tongue projection in A. tigrinum is the combined result of activity in four muscles: the geniohyoideus, Subarcualis rectus 1, intermandibularis posterior, and interhyoideus. Stimulation of the Subarcualis rectus 1 alone does not cause tongue projection. The kinetic model produced from the kinematic and stimulation data involves both a dorsal vector (the resultant of the Subarcualis rectus 1, intermandibularis posterior, and interhyoideus) and a ventral vector (the geniohyoideus muscle), which sum to produce a resultant anterior vector that directs tongue motion out of the mouth and toward the prey. This model generates numerous testable predictions about tongue function and provides a mechanistic basis for the hypothesis that tongue projection in salamanders evolved from primitive intraoral manipulative action of the hyobranchial apparatus.