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.     

Light microscopy and scanning electron microscopy studies on the reduction of the tongue microstructures in the white stork (Ciconia ciconia,Aves)

The structure of the tongue in the white stork (Ciconia ciconia) is observed macroscopically and under light and scanning electron microscopy. Our observations of the tongue reveal a rare terminal reduction of the size of the tongue and microstructures of the lingual mucosa among the investigations of birds published so far. The short, triangular tongue with a pointed tip is approximately 2.5 cm long in the adult and is situated in the caudal part of the oral cavity close to the laryngeal prominence. On the dorsal surface of the tongue, no typical mucosa microstructures like lingual papillae, median groove or lingual prominence are observed. The main structure of the tongue is composed of rostral part of hyoid apparatus, that is, entoglossal cartilage connects with basihyoid. Very thin mucosa is composed of fibrous connective tissue covered with orthokeratinized epithelium. No lingual glands and muscles are observed in the lamina propria of mucosa. Even though the triangular shape of the tongue in the white stork is typical for birds, the inner structure of the reduced organ is composed only of flat cartilagineous entoglossum of hyoid apparatus. During feeding behaviour of the white stork, the food transportation in oral cavity called cranio‐inertial transport is undoubtedly affected by structural reduction of the tongue.     

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.     

Mechanism of intraoral transport in macaques

All mammals have the same divisions of cyclic movement of tongue and hyoid during mastication: a protraction or forward phase that begins at minimum gape, and a retraction or return phase. Nonanthropoid mammals transport food from the oral cavity to the oropharynx during the return phase; food on the dorsal surface of the tongue moves distally while the tongue is retracted. Macaques, however, transport food during the protraction phase of tongue/hyoid movement. Food is squeezed posteriorly by contact between the tongue surface and the palate anterior to the food. This mechanism of transport is occasionally seen in nonanthropoid mammals when they are transporting liquids from the oral cavity to the oropharynx. It has, however, not been seen when these mammals transport solid food. One morphological basis for this difference is the reduction in height of the rugae of the palate of macaques. In most mammals these rugae are pronounced ridges that are able to hold food in place during protraction as the tongue slides forward beneath the food. Anthropoids and other mammals differ in the way they store food prior to swallowing. When macaques transport food to the oropharynx, usually they swallow in the next cycle, but always in the next 2 or 3 cycles. Most mammals transport and store food in the oropharynx for several cycles before a swallow clears that region of food. This behavior is correlated with differences in morphology of the oropharynx; anthropoids have reduced valleculae, the area in which other mammals store food prior to swallowing.     

The use of the tongue and hyoid apparatus during feeding in lizards (Ctenosaura similis and Tupinambis nigropunctatus)

The use of the tongue and hyoid is examined in cineradiographic and electromyographic investigations of feeding in two species of lizards, Ctenosaura similis (Iguanidae) and Tupinambis nigropunctatus (Teiidae). In both animals food is transported through the oral cavity by regular cycles of the tongue. Tongue movements correlate with jaw and hyoid movement. Similarities between the two animals in the use of the tongue in food transport, lapping, pharyngeal packing, and pharyngeal emptying are detailed. Mechanisms of tongue protrusion are examined and it is shown that the tongue in Tupinambis is relatively more protrusible than in Ctenosaura. This difference is complementary with data on the greater reliance of Tupinambis on the tongue as a sensory organ. Tupinambis further differs from Ctenosaura in possessing a greater mobility of the hyoid. In many features of tongue use in food transport, lizards resemble mammals, supporting postulations of a basic pattern of intra-oral food transport. However, whether this pattern can be attributed to convergence or a common, primitive neural pattern of control cannot be distinguished. Lizards lack two major characteristics of mammalian food transport: regular masticatory cycles and an internal swallowing mechanism.     

Effect of sensory input from the tongue on jaw movement in normal feeding in the opossum

Opossums were presented with solid and liquid foods. The movements of the jaw and tongue were recorded cineradiographically together with recordings of the EMG activity in muscles opening the jaw and moving the base of the tongue (hyoid). The jaw opening in each cycle was in two stages--01 and 02; 01 had a constant amplitude irrespective of the food ingested. Ingestion of liquid (which involved continuous accumulation of a liquid bolus in the valleculae prior to swallowing) was associated with cycles of oral movement in which 02 was small; tongue retraction was associated with this opening. In contrast, solid and semisolid food ingestion was associated with large angles of jaw opening in 02 that also coincided with the tongue retraction. In this latter case a characteristic pattern of EMG activity, in which all the muscles moving the hyoid were simultaneously active, was added to the pattern seen in lapping; this additional activity had an EMG pattern that was consistent with a jaw opening reflex. The findings contrast with other reports that the jaw opening reflex is suppressed in mastication. Experimentally induced tongue contact with a variety of solid surfaces during lapping (an activity involving accumulation of a liquid bolus in the valleculae) induced neither increased jaw opening nor the additional EMG pattern. However, in situations when there was no bolus in the valleculae, additional jaw opening activity was elicited when the tongue contracted solids intra- or extra-orally. It is suggested that the ability of sensory input, from the anterior tongue, to elicit a jaw opening reflex and to change the type of jaw/tongue cycle was dependent upon the extent of bolus accumulation in the valleculae and therefore indirectly upon the consistency of the food.     

A cineradiographic and electromyographic study of mastication in Tenrec ecaudatus

Regular chewing was studied in the specialized Malagasy insectivore Tenrec ecaudatus with the aid of precisely correlated electromyography of the main adductors, digastrics, and two hyoid muscles and cineradiography for which metallic markers were placed in the mandibles, tongue, and hyoid bone. During the power stroke the body of the mandible moves dorsally and medially. The medially directed component of movement at this time is greatly increased by simultaneous rotation of the mandible about its longitudinal axis. The highly mobile symphysis, spherical dentary condyle, loss of superficial masseter muscle and zygoma, and the simplified zalamnodont molars all appear to be related to the large amount of mandibular rotation that occurs during occlusion. The balancing side lateral pterygoid muscle (inferior head) apparently shifts the working side mandible laterally during the last part of opening and the first part of closing. The working side temporalis and the superficial masseter muscle are both responsible for the shift back to the midline. The temporalis is usually active to the same extent on the working and balancing sides during the power stroke. The level of activity (amplitude) of the temporalis and duration of the power stroke increase with harder foods. Whenever soft foods are chewed, the superficial masseter is only active on the working side; whenever foods of increasing hardness are chewed, its level of activity on the balancing side increases to approach that of the working side. Mandibular rotation is greatly reduced when hard foods are chewed.     

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.     

白鱀豚(齿鲸亚目、白鱀豚科)的肌学研究——Ⅱ.咀嚼肌、舌肌和舌骨肌、喉肌和咽肌

前已报道白鱀豚(Lipotes vexillifer)的皮肤肌、肩和鳍肢肌肉、胸壁肌肉、腹壁肌肉,颈、背和尾部肌肉(周开亚等,1981)。本篇包括咀嚼肌、舌和舌骨部肌肉、喉部肌肉及鼻咽和咽部肌肉。 一、咀嚼肌(图1、2) 咬肌m.masseter很薄,起自颧突后部的腹缘,颧突下方颞肌的筋膜及颧弓的腹缘。纤维尾腹向,止于下颌骨后部下半的外侧。后端有少量纤维绕过下颌骨腹缘,止于下颌内脂肪体,紧贴下颌骨的部分咬肌纤维已脂化形成下颌外脂肪体。咬肌仍有提起下颌的作用,但力量很弱。     

Feeding specializations of the Mexican burrowing toad, Rhinophrynus dorsalis (Anura: Rhinophrynidae)

Of the several, unrelated anuran taxa that feed underground, the Neotropical pipoid, Rhinophrynus dorsalis , seems to be the most specialized ant- and termite-feeder. The snout is covered with a curious and apparently unique epidermal armour. The buccal and oesophageal linings are ornately folded. The lips effect a double closure along the long, wedgeshaped, edentate maxillary arch. Peculiar submandibular glands seem to enhance the seal of the lips. The results of morphological, cinematographic, and muscle stimulation studies reveal that Rhinophrynus has a mechanism of tongue protrusion basically distinct from that of other frogs that project their tongues by means of a lingual flip. In Rhinophrynus , the intrinsic tongue muscles act to stiffen the organ, exerting hydrostatic pressure on the fluid contents of the lingual sinus. Actual protrusion of the tongue through the buccal groove involves shifting the organ forward via protraction of the hyoid by muscles extrinsic to the tongue—a mode that is unique among anurans and one highly suited for securing small insect prey in subterranean burrows.     

Differential sensitivity of tongue areas and palate to electrical stimulation: a suprathreshold cross-modal matching study

A cross-modal matching procedure was used, in twelve subjects,to evaluate regional differences in suprathreshold sensitivityof the oral cavity to electrogustometric stimulation. Stimulationof five loci on each side of the oral cavity was performed:tongue tip (one cm from the midline), anterior tongue side (2.5cm from tip on lateral margin), posterior tongue side (regionof the foliate papillae), posterior medial tongue (one cm frommidline on circumvallate papillae), and soft palate (one cmfrom midline, one cm above superior pole of anterior palatinearch). The tip of the tongue was significantly more sensitivethan the other areas to electric stimulation, as evidenced bythe slope and absolute position of the psychophysical powerfunctions. Strong correlations were observed in the sensitivitymeasures across tongue loci and between tongue and palate sides.No effects of subject gender or mouth side were found.     

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.     

Morphology and evolution of the ectethmoid-mandibular articulation in the meliphagidae (Aves)

The ectethmoid-mandibular articulation in Melithreptus and Manorina (Meliphagidae: Aves) consists of the dorsal mandibular process fitting into and abutting against the ventral ectethmoid fossa; it forms a brace for the mandible. This articulation in Melithreptus is a typical diarthrosis with long folded capsular walls. The mandible, thus, has two separate articulations, each with a different axis of rotation. No other genus of Meliphagidae (except Ptiloprora) or any other avian family possesses a similar feature. The jaw and tongue musculature of Melithreptus are described. The two muscles opening the jaws are well developed, while those closing the jaws are small. The tongue muscles show no special developments. A large maxillary gland, presumably muscus secreting, covers the ventral surface of the jaw muscles. Its duct opens into the oral cavity just behind the tip of the upper jaw. The frilled tip of the tongue rests against the duct opening. The ectethmoid-mandibular articulation braces the adducted mandible against dorsoposteriorly directed forces. The mandible can be held closed without a compression force exerted by the mandible on the quadrate, permitting the bird to raise its upper jaw with greater ease and less loss of force. The tongue can be protruded through the slight gap between the jaws, moving against the duct opening and thus be coated with mucus. Presumably, these birds capture insects with their sticky tongue. Hence, the ectethmoid-mandibular articulation is an adaptation for this feeding method; it evolved independently in three genera of the Meliphagidae. The ectethmoid-mandibular articulation demonstrates that a bone can have two articulations with different axes of rotation, that the two articular halves can separate widely, and that articular cartilages can be flat and remain in contact over a large area. Its function suggests that the basitemporal articulation of the mandible found in many other birds has a similar function. And it demonstrates that in the evolution of the mammalian dentary-squamosal articulation, the new hinge did not have to lie on the same rotational axis as the existing quadrate-articular hinge.     

Modulation of upper airway muscle activities by bronchopulmonary afferents.

Here we review the influence of bronchopulmonary receptors (slowly and rapidly adapting pulmonary stretch receptors, and pulmonary/bronchial C-fiber receptors) on respiratory-related motor output to upper airway muscles acting on the larynx, tongue, and hyoid arch. Review of the literature shows that all muscles in all three regions are profoundly inhibited by lung inflation, which excites slowly adapting pulmonary stretch receptors. This widespread coactivation includes the recruitment of muscles that have opposing mechanical actions, suggesting that the stiffness of upper airway muscles is highly regulated. A profound lack of information on the modulation of upper airway muscles by rapidly adapting receptors and bronchopulmonary C-fiber receptors prohibits formulation of a conclusive opinion as to their actions and underscores an urgent need for new studies in this area. The preponderance of the data support the view that discharge arising in slowly adapting pulmonary stretch receptors plays an important role in the initiation of the widespread and highly coordinated recruitment of laryngeal, tongue, and hyoid muscles during airway obstruction.     

Food transport through the anterior oral cavity in macaques

Intraoral transport, the movement of food or liquid through the oral cavity and oropharynx, is a major component of feeding behavior. Stage I transport, transport through the oral cavity prior to mastication, has been described for several mammals (Franks et al.: Arch. Oral Biol. 30:539, 1985; Hiiemae and Crompton: Hildebrand et al. (eds.): Functional Vertebrate Morphology, Cambridge, MA, Belknap Press, 1985). Previous work (Franks et al.: Am. J. Phys. Anthropol. 65:275, 1984) indicated that this was not a significant behavior in macaques in a laboratory setting, because food was ingested directly to the region of the cheek teeth. Although relatively infrequent in a captive situation, stage I transport does occur in long-tailed macaques through a mechanism similar to other mammals, but also subject to unique aspects of primate anatomy. Transport takes several cycles during which the food moves back and forth in an anterior/posterior direction, due to tongue movements. Because anthropoid primates lack the pronounced rugae that in other mammals prevent the anterior displacement of a bolus, stage I transport uses the rounded arch of the upper, anterior dentition to hold the food during the forward movement of the tongue. During the final cycle of transport, a pronounced twisting of the tongue, along a midline anteroposterior axis helps funnel the food item toward the postcanine teeth for subsequent mastication. This twisting, which was described in humans by Abd-El-Malek (J. Anat. 100:215, 1955) but not within the context of jaw movement, occurs prior to the closing phase of the jaw cycle.     

Respiratory function of hyoid muscles and hyoid arch

The position of the hyoid arch suggests that it supports soft tissue surrounding the upper airway (UA) and can act to maintain UA patency. We also suspected that muscles inserting on the hyoid arch might show respiratory patterns of activity that could be affected by respiratory stimuli. To test these possibilities, we moved the hyoid arch ventrally in six anesthetized dogs either by traction on it or by stimulation of hyoid muscles. UA resistance was decreased 73 +/- (SE) 6% and 72 +/- 6% by traction and stimulation during expiration and 57 +/- 15% and 52 +/- 8% during inspiration. Moving averages of the geniohyoid (GH) and thyrohyoid (TH) obtained in six other dogs breathing 100% O2 showed phasic respiratory activity while the sternohyoid (SH) showed phasic respiratory activity in only two of these animals and no activity in four. With progressive hypercapnia, GH and TH increased as did SH when activity was already present. Airway occlusion at end expiration augmented and prolonged inspiratory activity in the hyoid muscles but did not elicit SH activity if not already present. Occlusion at end inspiration suppressed phasic activity in hyoid muscles for as long as in the diaphragm. After vagotomy activity increased and became almost exclusively inspiratory. Activity appeared in SH when not previously present. Duration and amplitude of hyoid muscle activity were increased with negative UA pressure and augmented breaths. We conclude that the hyoid arch and muscles can strongly affect UA flow resistance. Hyoid muscles show responses to chemical, vagal, and negative pressure stimuli similar to other UA muscles.     

Twist and chew: three-dimensional tongue kinematics during chewing in macaque primates

Three-dimensional (3D) tongue movements are central to performance of feeding functions by mammals and other tetrapods, but 3D tongue kinematics during feeding are poorly understood. Tongue kinematics were recorded during grape chewing by macaque primates using biplanar videoradiography. Complex shape changes in the tongue during chewing are dominated by a combination of flexion in the tongue''s sagittal planes and roll about its long axis. As hypothesized for humans, in macaques during tongue retraction, the middle (molar region) of the tongue rolls to the chewing (working) side simultaneous with sagittal flexion, while the tongue tip flexes to the other (balancing) side. Twisting and flexion reach their maxima early in the fast close phase of chewing cycles, positioning the food bolus between the approaching teeth prior to the power stroke. Although 3D tongue kinematics undoubtedly vary with food type, the mechanical role of this movement—placing the food bolus on the post-canine teeth for breakdown—is likely to be a powerful constraint on tongue kinematics during this phase of the chewing cycle. The muscular drivers of these movements are likely to include a combination of intrinsic and extrinsic tongue muscles.     

The suction mechanism of the pipid frog,Pipa pipa (Linnaeus, 1758)

Most suction‐feeding, aquatic vertebrates create suction by rapidly enlarging the oral cavity and pharynx. Forceful enlargement of the pharynx is powered by longitudinal muscles that retract skeletal elements of the hyoid, more caudal branchial arches, and, in many fish, the pectoral girdle. This arrangement was thought to characterize all suction‐feeding vertebrates. However, it does not exist in the permanently aquatic, tongueless Pipa pipa , an Amazonian frog that can catch fish. Correlating high‐speed (250 and 500 fps) video records with anatomical analysis and functional tests shows that fundamental features of tetrapod body design are altered to allow P. pipa to suction‐feed. In P. pipa , the hyoid apparatus is not connected to the skull and is enclosed by the pectoral girdle. The major retractor of the hyoid apparatus arises not from the pectoral girdle but from the femur, which lies largely within the soft tissue boundaries of the trunk. Retraction of the hyoid is coupled with expansion of the anterior trunk, which occurs when the hypertrophied ventral pectoral elements are depressed and the urostyle and sacral vertebra are protracted and slide forward on the pelvic girdle, thereby elongating the entire trunk. We suggest that a single, robust pair of muscles adduct the cleithra to depress the ventral pectoral elements with force, while modified tail muscles slide the axial skeleton cranially on the pelvic girdle. Combined hyoid retraction, axial protraction, and pectoral depression expand the buccopharyngeal cavity to a volume potentially equal to that of the entire resting body of the frog. Pipa may be the only tetrapod vertebrate clade that enlarges its entire trunk during suction‐feeding.     

A control model of human tongue movements in speech

Tongue movements during speech production have been investigated by means of a simple yet realistic biomechanical model, based on a finite elements modeling of soft tissues, in the framework of the equilibrium point hypothesis (-model) of motor control. In particular, the model has been applied to the estimation of the “central” control commands issued to the muscles, for a data set of mid-sagittal digitized tracings of vocal tract shape, r ecorded by means of low-intensity X-ray cineradiographies during speech. In spite of the highly non-linear mapping between the shape of the oral cavity and its acoustic consequences, the organization of control commands preserves the peculiar spatial organization of vowel phonemes in acoustic space. A factor analysis of control commands, which have been decomposed into independent or “orthogonal” muscle groups, has shown that, in spite of the great mobility of the tongue and the highly complex arrangement of tongue muscles, its movements can be explained in terms of the activation of a small number of independent muscle groups, each corresponding to an elementary or “primitive” movement. These results are consistent with the hypothesis that the tongue is controlled by a small number of independent “articulators”, for which a precise biomechanical substrate is provided. The influence of the effect of jaw and hyoid movements on tongue equilibrium has also bee n evaluated, suggesting that the bony structures cannot be considered as a moving frame of reference, but, indeed, there may be a substantial interaction between them and the tongue, that may only be accounted for by a “global” model. The reported results also define a simple control model for the tongue and, in analogy with similar modelling studies, they suggest that, because of the peculiar geometrical arrangement of tongue muscles, the central nervous system (CNS) may not need a de tailed representation of tongue mechanics but rather may make use of a relatively small number of muscle synergies, that are invariant over the whole space of tongue configurations. Received: 27 August 1996 / Accepted in revised form: 25 February 1997     

Prey capture in the lizard Agama stellio

Prey capture in Agama stellio was recorded by high-speed video in combination with the electrical activity of both jaw and hyolingual muscles. Quantification of kinematics and muscle activity patterns facilitated their correlation during kinematic phases. Changes in angular velocity of the gape let the strike be subdivided into four kinematic phases: slow open (SOI and SOII), fast open (FO), fast close (FC), and slow close-power stroke (SC/PS). The SOI phase is marked by initial activity in the tongue protractor, the hyoid protractor, and the ring muscle. These muscles project the tongue beyond the anterior margin of the jaw. During the SOII phase, a low level of activity in the jaw closers correlates with a decline of the jaw-opening velocity. Next, bilateral activity in the jaw openers defines the start of the FO phase. This activity ends at maximal gape. Simultaneously, the hyoid retractor and the hyoglossus become active, causing tongue retraction during the FO phase. At maximal gape, the jaw closers contract simultaneously, initiating the FC phase. After a short pause, they contract again and the prey is crushed during the SC/PS phase. Our results support the hypothesis of tongue projection in agamids by Smith ([1988] J. Morphol. 196:157–171), and show some striking similarities with muscle activity patterns during the strike in chameleons (Wainwright and Bennett [1992a] J. Exp. Biol. 168:1–21). Differences are in the activation pattern of the hyoglossus. The agamid tongue projection mechanism appears to be an ideal mechanical precursor for the ballistic tongue projection mechanism of chameleonids; the key derived feature in the chameleon tongue projection mechanism most likely lies in the changed motor pattern controlling the hyoglossus muscle. © 1995 Wiley-Liss, Inc.