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.     

Reconstruction of cranial and hyobranchial muscles in the triassic temnospondyl Gerrothorax provides evidence for akinetic suction feeding

The cranial and hyobranchial muscles of the Triassic temnospondyl Gerrothorax have been reconstructed based on direct evidence (spatial limitations, ossified muscle insertion sites on skull, mandible, and hyobranchium) and on phylogenetic reasoning (with extant basal actinopterygians and caudates as bracketing taxa). The skeletal and soft‐anatomical data allow the reconstruction of the feeding strike of this bottom‐dwelling, aquatic temnospondyl. The orientation of the muscle scars on the postglenoid area of the mandible indicates that the depressor mandibulae was indeed used for lowering the mandible and not to raise the skull as supposed previously and implies that the skull including the mandible must have been lifted off the ground during prey capture. It can thus be assumed that Gerrothorax raised the head toward the prey with the jaws still closed. Analogous to the bracketing taxa, subsequent mouth opening was caused by action of the strong epaxial muscles (further elevation of the head) and the depressor mandibulae and rectus cervicis (lowering of the mandible). During mouth opening, the action of the rectus cervicis muscle also rotated the hyobranchial apparatus ventrally and caudally, thus expanding the buccal cavity and causing the inflow of water with the prey through the mouth opening. The strongly developed depressor mandibulae and rectus cervicis, and the well ossified, large quadrate‐articular joint suggest that this action occurred rapidly and that powerful suction was generated. Also, the jaw adductors were well developed and enabled a rapid mouth closure. In contrast to extant caudate larvae and most extant actinopterygians (teleosts), no cranial kinesis was possible in the Gerrothorax skull, and therefore suction feeding was not as elaborate as in these extant forms. This reconstruction may guide future studies of feeding in extinct aquatic tetrapods with ossified hyobranchial apparatus. J. Morphol., 2013. © 2012 Wiley Periodicals, Inc.     

Functional analysis of a specialized prey processing behavior: winnowing by surfperches (Teleostei: Embiotocidae).

Several surfperches (Embiotocidae), including the black surfperch, Embiotoca jacksoni, exhibit a specialized prey handling behavior known as winnowing, in which ingested food and non-nutritive debris are separated within the oropharyngeal cavity. Prey items are swallowed, and unpalatable material is ejected from the mouth. Winnowing is believed to play an important role in the partitioning of food resources among sympatric embiotocids. We present a mechanistic model for this separative prey processing based on high-speed video analysis, cineradiography, electromyography, and buccal and opercular cavity pressure transducer recording. Winnowing by embiotocids is characterized by premaxillary protrusions repeated cyclically with reduced oral gape. Protrusion is accompanied by depression of the hyoid apparatus and adduction of the opercula. Alternating expansion and contraction of the buccal and opercular cavities generate regular pressure waveforms that indicate bidirectional water flow during processing. Separation of food from debris by Embiotoca jacksoni occurs in three phases. The prey-debris bolus is transported anteriorly and posteriorly within the oropharyngeal cavity and is then sheared by the pharyngeal jaws. Mechanical processing is complemented by the rinsing action of water currents during hydraulic prey transport. The feeding apparatus of Embiotoca jacksoni is functionally versatile, although not obviously specialized relative to that of nonwinnowing surfperches. Protrusion of the premaxillae and depression of the hyoid apparatus are critical to both prey capture and subsequent prey processing. The pharyngeal jaws exhibit kinematic patterns during separation of food from debris distinct from those observed during mastication of uncontaminated prey. This behavioral flexibility facilitates resource partitioning and the coexistence of E. jacksoni in sympatric embiotocid assemblages.     

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.     

Evolution of the feeding mechanism in primitive actionopterygian fishes: A functional anatomical analysis of Polypterus,Lepisosteus, and Amia

The comparative functional anatomy of feeding in Polypterus senegalus, Lepisosteus oculatus, and Amia calva, three primitive actinopterygian fishes, was studied by high-speed cinematography (200 frames per second) synchronized with electromyographic recordings of cranial muscle activity. Several characters of the feeding mechanism have been identified as primitive for actinopterygian fishes: (1) Mandibular depression is mediated by the sternohyoideus muscle via the hyoid apparatus and mandibulohyoid ligament. (2) The obliquus inferioris and sternohyoideus muscles exhibit synchronous activity at the onset of the expansive phase of jaw movement. (3) Activity in the adductor operculi occurs in a double burst pattern—an initial burst at the onset of the expansive phase, followed by a burst after the jaws have closed. (4) A median septum divides the sternohyoideus muscle into right and left halves which are asymmetrically active during chewing and manipulation of prey. (5) Peak hyoid depression occurs only after peak gape has been reached and the hyoid apparatus remains depressed after the jaws have closed. (6) The neurocranium is elevated by the epaxial muscles during the expansive phase. (7) The adductor mandibulae complex is divided into three major sections—an anterior (suborbital) division, a medial division, and a posterolateral division. In Polypterus, the initial strike lasts from 60 to 125 msec, and no temporal overlap in muscle activity occurs between muscles active at the onset of the expansive phase (sternohyoideus, obliquus superioris, epaxial muscles) and the jaw adductors of the compressive phase. In Lepisosteus, the strike is extremely rapid, often occuring in as little as 20 msec. All cranial muscles become active within 10 msec of each other, and there is extensive overlap in muscle activity periods. Two biomechanically independent mechanisms mediate mandibular depression in Amia, and this duality in mouth-opening couplings is a shared feature of the halecostome fishes. Mandibular depression by hyoid retraction, and intermandibular musculature, consisting of an intermandibularis posterior and interhyoideus, are hypothesized to be primitive for the Teleostomi.     

Patterns of Evolution in the Feeding Mechanism of Actinopterygian Fishes

SYNOPSIS. Structural and functional patterns in the evolutionof the actinopterygian feeding mechanism are discussed in thecontext of the major monophyletic lineages of ray-finned fishes.A tripartite adductor mandibulae contained in a maxillary-palatoquadratechamber and a single mechanism of mandibular depression mediatedby the obliquus inferioris, sternohyoideus, and hyoid apparatusare primitive features of the Actinopterygii. Halecostome fishesare characterized by having an additional mechanism of mandibulardepression, the levator operculi—opercular series coupling,and a maxilla which swings anteriorly during prey capture. Theseinnovations provide the basis for feeding by inertial suctionwhich is the dominant mode of prey capture throughout the halecostomeradiation. A remarkably consistent kinematic profile occursin all suction-feeding halecostomes. Teleost fishes possessa number of specializations in the front jaws including a geniohyoideusmuscle, loss of the primitive suborbital adductor component,and a mobile premaxilla. Structural innovations in teleost pharyngealjaws include fusion of the dermal tooth plates with endoskeletalgill arch elements, the occurrence of a pharyngeal retractormuscle, and a shift in the origin of the pharyngohyoideus. Thesespecializations relate to increased functional versatility ofthe pharyngeal jaw apparatus as demonstrated by an electromyographicstudy of pharyngeal muscle activity in Esox and Ambloplites.The major feature of the evolution of the actinopterygian feedingmechanism is the increase in structural complexity in both thepharyngeal and front jaws. Structural diversification is a functionof the number of independent biomechanical pathways governingmovement.     

Functional morphology of the feeding apparatus,feeding constraints,and suction performance in the nurse shark Ginglymostoma cirratum

The nurse shark, Ginglymostoma cirratum, is an obligate suction feeder that preys on benthic invertebrates and fish. Its cranial morphology exhibits a suite of structural and functional modifications that facilitate this mode of prey capture. During suction‐feeding, subambient pressure is generated by the ventral expansion of the hyoid apparatus and the floor of its buccopharyngeal cavity. As in suction‐feeding bony fishes, the nurse shark exhibits expansive, compressive, and recovery kinematic phases that produce posterior‐directed water flow through the buccopharyngeal cavity. However, there is generally neither a preparatory phase nor cranial elevation. Suction is generated by the rapid depression of the buccopharyngeal floor by the coracoarcualis, coracohyoideus, and coracobranchiales muscles. Because the hyoid arch of G. cirratum is loosely connected to the mandible, contraction of the rectus cervicis muscle group can greatly depress the floor of the buccopharyngeal cavity below the depressed mandible, resulting in large volumetric expansion. Suction pressures in the nurse shark vary greatly, but include the greatest subambient pressures reported for an aquatic‐feeding vertebrate. Maximum suction pressure does not appear to be related to shark size, but is correlated with the rate of buccopharyngeal expansion. As in suction‐feeding bony fishes, suction in the nurse shark is only effective within approximately 3 cm in front of the mouth. The foraging behavior of this shark is most likely constrained to ambushing or stalking due to the exponential decay of effective suction in front of the mouth. Prey capture may be facilitated by foraging within reef confines and close to the substrate, which can enhance the effective suction distance, or by foraging at night when it can more closely approach prey. J. Morphol., 2008. © 2008 Wiley‐Liss, Inc.     

Feeding in Atractaspis (Serpentes: Atractaspididae): a study in conflicting functional constraints

African fossorial colubroid snakes of the genus Atractaspis have relatively long fangs on short maxillae, a gap separating the pterygoid and palatine bones, a toothless pterygoid, and a snout tightly attached to the rest of the skull. They envenomate prey with a unilateral backward stab of one fang projected from a closed mouth. We combined structural reanalysis of the feeding apparatus, video records of prey envenomation and transport, and manipulations of live and dead Atractaspis to determine how structure relates to function in this unusual genus of snakes. Unilateral fang use in Atractaspis is similar to unilateral slashing envenomation by some rear-fanged snakes, but Atractaspis show no maxillary movement during prey transport. Loss of pterygoid teeth and maxillary movement during transport resulted in the inability to perform. 'pterygoid walk' prey transport. Atractaspis transport prey through the oral cavity using movement cycles in which mandibular adduction, anterior trunk compression, and ventral flexion of the head alternate with mandibular abduction and extension of head and anterior trunk over the prey. Inefficiencies in manipulation and early transport of prey are offset by adaptability of the envenomating system to various prey types in both enclosed and open spaces and by selection of prey that occupy burrows or tunnels in soil. Atractaspis appears to represent the evolutionary endpoint of a functional conflict between envenomation and transport in which a rear-fanged envenomating system has been optimized at the expense of most, if not all, palatomaxillary transport function.     

Morphological and functional bases of durophagy in the queen triggerfish,Balistes vetula (Pisces,tetraodontiformes)

Tetraodontiform fishes are characterized by jaws specialized for powerful biting and a diet dominated by hard-shelled prey. Strong biting by the oral jaws is an unusual feature among teleosts. We present a functional morphological analysis of the feeding mechanism of a representative tetraodontiform, Balistes vetula. As is typical for the order, long, sharp, strong teeth are mounted on the short, robust jaw bones of B. vetula. The neurocranium and suspensorium are enlarged and strengthened to serve as sites of attachment for the greatly hypertrophied adductor mandibulae muscles. Electromyographic recordings made from 11 cranial muscles during feeding revealed four distinct behaviors in the feeding repertoire of B. vetula. Suction is used effectively to capture soft prey and is associated with a motor pattern similar to that reported for many other teleosts. However, when feeding on hard prey, B. vetula directly bit the prey, exhibiting a motor pattern very different from that of suction feeding. During buccal manipulation, repeated cycles of jaw opening and closing (biting) were coupled with rapid movement of the prey in and out of the mouth. Muscle activity during buccal manipulation was similar to that seen during bite-captures. A blowing behavior was periodically employed during prey handling, as prey were forcefully “spit out” from the mouth, either to reposition them or to separate unwanted material from flesh. The motor pattern used during blowing was distinct from similar behaviors described for other fishes, indicating that this behaviors may be unique to tetraodontiforms. Thus B. vetula combines primitive behaviors and motor patterns (suction feeding and buccal manipulation) with specialized morphology (strong teeth, robust jaws, and hypertrophied adductor muscles) and a novel behavior (blowing) to exploit armored prey such as sea urchins molluscs, and crabs. © 1993 Wiley-Liss, Inc.     

A functional morphospace for the skull of labrid fishes: patterns of diversity in a complex biomechanical system

The Labridae (including wrasses, the Odacidae and the Scaridae) is a species‐rich group of perciform fishes whose members are prominent inhabitants of warm‐temperate and tropical reefs worldwide. We analyse functionally relevant morphometrics for the feeding apparatus of 130 labrid species found on the Great Barrier Reef and use these data to explore the morphological and mechanical basis of trophic diversity found in this assemblage. Morphological measurements were made that characterize the functional and mechanical properties of the oral jaws that are used in prey capture and handling, the hyoid apparatus that is used in expanding the buccal cavity during suction feeding, and the pharyngeal jaw apparatus that is used in breaking through the defences of shelled prey, winnowing edible matter from sand and other debris, and pulverizing the algae, detritus and rock mixture eaten by scarids (parrotfishes). A Principal Components Analysis on the correlation matrix of a reduced set of ten variables revealed complete separation of scarids from wrasses on the basis of the former having a small mouth with limited jaw protrusion, high mechanical advantage in jaw closing, and a small sternohyoideus muscle and high kinematic transmission in the hyoid four‐bar linkage. Some scarids also exhibit a novel four‐bar linkage conformation in the oral jaw apparatus. Within wrasses a striking lack of strong associations was found among the mechanical elements of the feeding apparatus. These weak associations resulted in a highly diverse system in which functional properties occur in many different combinations and reflect variation in feeding ecology. Among putatively monophyletic groups of labrids, the cheilines showed the highest functional diversity and scarids were moderately diverse, in spite of their reputation for being trophically monomorphic and specialized. We hypothesize that the functional and ecological diversity of labrids is due in part to a history of decoupled evolution of major components of the feeding system (i.e. oral jaws, hyoid and pharyngeal jaw apparatus) as well as among the muscular and skeletal elements of each component. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 82 , 1–25.     

On the geometry and mechanics of tooth position in the white shark,Carcharodon carcharias

The teeth of captured specimens, of prepared museum specimens, and of high-speed videotape images of the white shark, Carcharodon carcharias, were compared with respect to (1) deviation of each tooth from the animal's midline and (2) the crown angle of the functional teeth along the jaw margin. Tooth position was measured either directly using a meter stick apparatus or derived from tracings of the video footage. Tooth positions were not statistically unique in any region of the upper or lower jaw but demonstrated less variability in crown angle within 30° of the midline (71.48° ± 10°). Videotape analysis of feeding sharks indicated an 8.7° increase in crown angle of the centermost teeth during bites where the jaws were closed through an angle of 20–35° and a 15.7° reduction in this same parameter during jaw adduction through 35° or more. Such changes in tooth orientation (relative to the rear of the buccal cavity) are ascribed to flexure of the cartilaginous jaws and cranium by the cranial musculature and possibly also to sliding of the tooth bed over the jaw. Outward rotation of the teeth and jaw rami describes a plucking action during feeding or prey sampling, while larger bites rotate the frontmost teeth inward towards the gullet. Functionally, this may make the teeth more effective at grasping small prey items or gouging chunks from larger prey. However, testing of the load required to remove teeth showed no significant increase in tensile resistance with reduced crown angle. © 1995 Wiley-Liss, Inc.     

Form and function of the feeding apparatus of Alligator mississippiensis

The architecture of the jaw muscles and their tendons of Alligator mississippiensis is described and their function examined by electromyography. Alligator grabs its prey with forward lunges or rapid lateral movements of the head. It does not engage in regular masticatory cycles. Prey is manipulated by inertial movements and the tongue does not appear to play any role in transport. The Mm. adductor mandibulae externus, adductor mandibulae posterior, and pterygoideus activate bilaterally and simultaneously during rapid closing or crushing. The M. pterygoideus does not act during prey holding whereas the Mm. adductor mandibulae externus, adductor mandibulae posterior continue to be active. The Mm. depressor mandibulae and intramandibularis are variably active during both jaw opening and closing.     

Functional morphology of tongue projection in Taricha torosa (Urodela: Salamandridae)

The kinematics of tongue projection by terrestrial adult California newts, Taricha torosa (Rathke, 1833), are described based upon high-speed cinematography. Tongue projection results from coupled anterior movements of the ceratohyals and branchial arches. Four distinct periods are defined during a projection sequence: preparation, tongue projection, tongue recovery and mouth closing. Key anatomical correlates of projection are described, with special emphasis on the mobility of the hyoid arch. Adult Taricha (Gray, 1850) have lost the mandibulo-hyoid ligament and reduced additional connective tissues present in larvae. These changes decouple the hyoid arch from mouth opening and release the ceratohyals to participate in a tongue projection system distinct from those of ambystomatids and plethodontids. These phylogenetic differences pose questions about the evolution of tongue projection systems in terrestrial urodeles. Currently available data are consistent with the interpretation that terrestrial urodeles have independently evolved specialized tongue projection systems at least twice: the ceratohyal-stable mode of plethodontids and the ceratohyal-mobile system of newts. In all cases, an essential underlying (= plesiomorphic) feature is the presence of the depressor mandibulae muscle. We regard this pathway for mouth opening as a prerequisite which liberated the hyobranchium for alternative function. Comprehensive studies of the mandibulo-hyoid ligament and depressor mandibulae will be vital to modelling the evolution of specialized tongue projection systems of urodeles.     

Morphological and videofluorographic study of the hyoid apparatus and its function in the rabbit (Oryctolagus cuniculus)

The anatomy of the hyoid apparatus and positional changes of the hyoid bone during mastication and deglutition are described in the New Zealand White rabbit (Oryctolagus cuniculus). A testable model is constructed to predict the range of movement during function of the hyoid, a bone entirely suspended by soft tissue. Frame-by-frame analysis of a videofluorographic tape confirms the accuracy of the prediction through observation of hyoid bone excursion during oral behavior. During chewing, translation of the hyoid bone is diminutive and irregular, lacking a clearly discernible path of excursion. However, some movements of the hyoid occur with regularity. During fast opening, anterodorsal movement of the hyoid is interrupted with an abrupt posteroventral depression when the bolus is moved posteriorly toward the cheek teeth by the tongue. This clockwise rotation (when viewed from the right side) of the hyoid accompanies jaw opening and is reversed (posteroventral movement) for the jaw closing sequence. Lateral movements of the hyoid may be slightly coupled to mandibular rotation in the horizontal plane. The findings suggest that the hyoid bone maintains a relatively static position during the dynamics of chewing. The primary function would be to provide a stable base for the movements of the tongue. Another possible function would be to control the position of the larynx within the pharyngeal cavity. Some characteristic features of the rabbit hyoid apparatus may be consequential to relatively erect posture and a saltatory mode of locomotion.     

A fish that uses its hydrodynamic tongue to feed on land

To capture and swallow food on land, a sticky tongue supported by the hyoid and gill arch skeleton has evolved in land vertebrates from aquatic ancestors that used mouth-cavity-expanding actions of the hyoid to suck food into the mouth. However, the evolutionary pathway bridging this drastic shift in feeding mechanism and associated hyoid motions remains unknown. Modern fish that feed on land may help to unravel the physical constraints and biomechanical solutions that led to terrestrialization of fish-feeding systems. Here, we show that the mudskipper emerges onto land with its mouth cavity filled with water, which it uses as a protruding and retracting ‘hydrodynamic tongue’ during the initial capture and subsequent intra-oral transport of food. Our analyses link this hydrodynamic action of the intra-oral water to a sequence of compressive and expansive cranial motions that diverge from the general pattern known for suction feeding in fishes. However, the hyoid motion pattern showed a remarkable resemblance to newts during tongue prehension. Consequently, although alternative scenarios cannot be excluded, hydrodynamic tongue usage may be a transitional step onto which the evolution of adhesive mucosa and intrinsic lingual muscles can be added to gain further independence from water for terrestrial foraging.     

Prey Transport Mechanisms in Blindsnakes and the Evolution of Unilateral Feeding Systems in Snakes

Most snakes ingest and transport their prey via a jaw ratchetingmechanism in which the left and right upper jaw arches are advancedover the prey in an alternating, unilateral fashion. This unilateraljaw ratcheting mechanism differs greatly from the hyolingualand inertial transport mechanisms used by lizards, both of whichare characterized by bilaterally synchronous jaw movements.Given the well-corroborated phylogenetic hypothesis that snakesare derived from lizards, this suggests that major changes occurredin both the morphology and motor control of the feeding apparatusduring the early evolution of snakes. However, most previousstudies of the evolution of unilateral feeding mechanisms insnakes have focused almost exclusively on the morphology ofthe jaw apparatus because there have been very few direct observationsof feeding behavior in basal snakes. In this paper I describethe prey transport mechanisms used by representatives of twofamilies of basal snakes, Leptotyphlopidae and Typhlopidae.In Leptotyphlopidae, a mandibular raking mechanism is used,in which bilaterally synchronous flexions of the lower jaw serveto ratchet prey into and through the mouth. In Typhlopidae,a maxillary raking mechanism is used, in which asynchronousratcheting movements of the highly mobile upper jaws are usedto drag prey through the oral cavity. These findings suggestthat the unilateral feeding mechanisms that characterize themajority of living snakes were not present primitively in Serpentes,but arose subsequently to the basal divergence between Scolecophidiaand Alethinophidia.     

Functional morphology of the feeding system in the ruff — Gymnocephalus cernua (L. 1758) — (Teleostei,Percidae)

The ruff, Gymnocephalus cernua, is a European freshwater fish that feeds by sucking up small invertebrates from the bottom of ponds and slow flowing rivers. The feeding movements have been studied by simultaneous electromyography of seventeen muscles of the head and cinematographic techniques. A theoretical model of movements imposes the functional demands of suction upon an abstraction of the form of a teleost head. Three phases in the feeding act, a preparatory phase, a suction phase and a transport phase, could be correlated with the observed movements and EMGs. Differences between the predicted and the actual movement are discussed. Two different types of feeding occur. The direction, magnitude and duration of the suction forces during feeding are modified, according to the position of the prey. A mechanism preventing early mandibular depression allows sudden and strong suction. Retardation of the suspensorial abduction during the overall expansion of the buccal cavity is ascribed to kinetic interrelations with the hyoid arch. Protrusion of the upper jaws also permits an earlier closure of the mouth and directs the food-containing waterflow posteriorly. When the fish is feeding on sinking prey, protrusion occurs later in the sequence of movements than when it is feeding from the bottom. As the protruded jaws produce a downwardly pointed mouth this retardation aims the suction force.     

Odontocete suction feeding: Experimental analysis of water flow and head shape

The role of cranial morphology in the generation of intraoral and oropharyngeal suction pressures in odontocetes was investigated by manipulating the jaw and hyolingual apparatus of submerged heads of three species presenting varied shapes. Hyoid and gular muscles were manually employed to depress and retract the tongue. Pressures were recorded at three locations in the oral cavity, as gape and site, speed, and force of pull were varied. A biomechanical model was also developed to evaluate pressure data. The species with the shortest, bluntest head and smallest mouth opening generated greater negative pressures. Suction generation diminished sharply as gape increased. Greatest negative pressures attained were around -45 mmHg (-6,000 Pa), a magnitude deemed suitable for capture of small live prey. Odontocetes utilizing this bidirectional flow system should profit by evolution of a rounder mouth opening through progressive shortening and widening of the rostrum and jaws, a trend evident in cranial measurements from fossil and recent odontocetes. Blunt heads correlate with anatomical, ecological, and behavioral traits associated with suction feeding. Small-gape suction (with minimally opened jaws) could be used by odontocetes of all head and oral shapes to draw prey sufficiently close to the mouth for suction ingestion or grasping via dentition. Principal limitations of the experimental and mathematical simulations include assumption of a stationary odontocete with static (open or closed) jaws and potential scaling issues with differently sized heads and gapes.     

Eating with a saw for a jaw: Functional morphology of the jaws and tooth‐whorl in Helicoprion davisii

The recent reexamination of a tooth‐whorl fossil of Helicoprion containing intact jaws shows that the symphyseal tooth‐whorl occupies the entire length of Meckel's cartilage. Here, we use the morphology of the jaws and tooth‐whorl to reconstruct the jaw musculature and develop a biomechanical model of the feeding mechanism in these early Permian predators. The jaw muscles may have generated large bite‐forces; however, the mechanics of the jaws and whorl suggest that Helicoprion was better equipped for feeding on soft‐bodied prey. Hard shelled prey would tend to slip anteriorly from the closing jaws due to the curvature of the tooth‐whorl, lack of cuspate teeth on the palatoquadrate (PQ), and resistance of the prey. When feeding on soft‐bodied prey, deformation of the prey traps prey tissue between the two halves of the PQ and the whorl. The curvature of the tooth‐whorl and position of the exposed teeth relative to the jaw joint results in multiple tooth functions from anterior to posterior tooth that aid in feeding on soft‐bodied prey. Posterior teeth cut and push prey deeper into the oral cavity, while middle teeth pierce and cut, and anterior teeth hook and drag more of the prey into the mouth. Furthermore, the anterior‐posterior edges of the teeth facilitate prey cutting with jaw closure and jaw depression. The paths traveled by each tooth during jaw depression are reminiscent of curved pathways used with slashing weaponry such as swords and knifes. Thus, the jaws and tooth‐whorl may have formed a multifunctional tool for capturing, processing, and transporting prey by cyclic opening and closing of the lower jaw in a sawing fashion. J. Morphol. 276:47–64, 2015. © 2014 Wiley Periodicals, Inc.