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A Herrel

Publications and source records attributed to A Herrel.

12 recordsLinked to original sources

Head shape and bite performance in xenosaurid lizards.

Bite performance in lizards influences many aspects of the animal's lifestyle. During feeding, inter- and intrasexual interactions, and defensive behavior, the ability to bite hard might be advantageous. Although biomechanical considerations predict clear relations between head shape and bite performance, this has rarely been tested. Here we investigate the effect of head shape on bite performance in three closely related species of xenosaurid lizards. Our data show that in this family of lizards, bite performance is mainly determined by head height, with high headed animals biting harder than flat headed ones. Species clearly differ in head shape and bite performance and show a marked sexual dimorphism. The dimorphism in head shape also results in an intersexual difference in bite performance. As head height is the major determinant of bite performance in xenosaurid lizards, trade offs between a crevice dwelling life-style and bite performance seem to occur. The evolutionary implications of these results are discussed. J. Exp. Zool. 290:101-107, 2001.

Animals↗

Morphology and histochemistry of the hyolingual apparatus in chameleons.

We reexamined the morphological and functional properties of the hyoid, the tongue pad, and hyolingual musculature in chameleons. Dissections and histological sections indicated the presence of five distinctly individualized pairs of intrinsic tongue muscles. An analysis of the histochemical properties of the system revealed only two fiber types in the hyolingual muscles: fast glycolytic and fast oxidative glycolytic fibers. In accordance with this observation, motor-endplate staining showed that all endplates are of the en-plaque type. All muscles show relatively short fibers and large numbers of motor endplates, indicating a large potential for fine muscular control. The connective tissue sheet surrounding the entoglossal process contains elastin fibers at its periphery, allowing for elastic recoil of the hyolingual system after prey capture. The connective tissue sheets surrounding the m. accelerator and m. hyoglossus were examined under polarized light. The collagen fibers in the accelerator epimysium are configured in a crossed helical array that will facilitate limited muscle elongation. The microstructure of the tongue pad as revealed by SEM showed decreased adhesive properties, indicating a change in the prey prehension mechanics in chameleons compared to agamid or iguanid lizards. These findings provide the basis for further experimental analysis of the hyolingual system.

Acetylcholinesterase↗

Functional implications of supercontracting muscle in the chameleon tongue retractors.

Chameleons capture prey items using a ballistic tongue projection mechanism that is unique among lizards. During prey capture, the tongue can be projected up to two full body lengths and may extend up to 600 % of its resting length. Being ambush predators, chameleons eat infrequently and take relatively large prey. The extreme tongue elongation (sixfold) and the need to be able to retract fairly heavy prey at any given distance from the mouth are likely to place constraints on the tongue retractor muscles. The data examined here show that in vivo retractor force production is almost constant for a wide range of projection distances. An examination of muscle physiology and of the ultrastructure of the tongue retractor muscle shows that this is the result (i) of active hyoid retraction, (ii) of large muscle filament overlap at maximal tongue extension and (iii) of the supercontractile properties of the tongue retractor muscles. We suggest that the chameleon tongue retractor muscles may have evolved supercontractile properties to enable a substantial force to be produced over a wide range of tongue projection distances. This enables chameleons successfully to retract even large prey from a variety of distances in their complex three-dimensional habitat.

Animals↗

Spatio-temporal gait characteristics of level and vertical locomotion in a ground-dwelling and a climbing gecko.

The effects of incline (vertical versus horizontal) on spatio-temporal gait characteristics (stride and step length, frequency, duty factor, degree of sprawling) were measured over a range of speeds in a ground-dwelling (Eublepharis macularius) and a climbing (Gekko gecko) species of gecko. Surprisingly, the climbing species also performs very well when moving on the horizontal substratum. In the present experiments, climbing speeds ranged from 0.6 to 1.2 m s(-1), whereas speeds for level locomotion were between 0.6 and 1.8 m s(-1). In contrast, the vertical climbing capacities of the ground-dweller are limited (speeds below 0.1 m s(-1 )versus level speeds between 0.2 and 1.1 m s(-1)). In general, we demonstrate that very little adjustment in gait characteristics is made by either species when they are forced to move on their non-habitual substratum. Moreover, gait characteristics differ little between the species despite the clear differences in ecological niche. Higher level or climbing speeds are realized mainly (or exclusively in the case of level locomotion in G. gecko) by increasing stride frequency. Stride lengths and duty factors vary with speed in the ground-dweller, but not in the climbing species. Step length and the degree of sprawling are speed-independent (except for hind-limb sprawling in G. gecko on the level). It is argued that this common strategy suits climbing (fixed spatial variables, no floating phases) rather than level locomotion.

Adaptation, Physiological↗

The mechanics of prey prehension in chameleons.

Iguanian lizards generally use their tongue to capture prey. Because lingual prehension is based on surface phenomena (wet adhesion, interlocking), the maximal prey size that can be captured is small. However, published records show that prey items eaten by chameleons include small vertebrates such as lizards and birds, indicating that these lizards are using a different prey prehension mechanism. Using high-speed video recordings, cineradiography, electromyography, nerve transection and stimulation experiments, we investigated the function of the tongue during prey capture. The results of these experiments indicate that chameleons have modified the primitive iguanian system by including a suction component in their prehension mechanism. Suction is generated by the activity of two modified intrinsic tongue muscles that pull the tongue pad inwards. Moreover, we demonstrate that the mechanism described here is a prerequisite for successful feeding.

Animals↗

Cranial kinesis in geckoes: functional implications.

Although it is generally assumed that cranial kinesis is a plesiomorphic characteristic in squamates, experimental data tend to contradict this hypothesis. In particular, coupled kinesis (i.e. streptostyly and mesokinesis) presumably arose independently in only a limited number of highly specialised groups. In this study, we investigated cranial kinesis in one of the most specialised of these groups: geckoes. On the basis of cineradiographic and electromyographic data, the fast opening and the slow closing/power stroke phases were modelled to elucidate possible functions of the observed kinesis. The results of these analyses show that the retraction of the muzzle unit during crushing is a self-reinforcing system that increases bite force and reduces the joint forces; the active protraction of the kinetic system during jaw opening, in contrast, enhances opening speed through the coupling of the intracranial units. It can be argued that cranial kinesis in geckoes is probably not an adaptive trait as such but, instead, a consequence of the 'Bauplan' of the cranial system in these animals. Presumably as a result of constructional constraints on the size of the jaw musculature and eyes, the supratemporal and postorbital bars were lost, which resulted in enormous mobility in the skull. To counteract the potential negative factors associated with this (decrease in bite force, skull damage), the kinetic system may have become coupled, and thus functional.

Adaptation, Physiological↗

Morphology of the feeding system in agamid lizards: ecological correlates.

The interaction of organismal design with ecology, and its evolutionary development are the subject of many functional and ecomorphological studies. Many studies have shown that the morphology and mechanics of the masticatory apparatus in mammals are adapted to diet. To investigate the relations between diet and the morphological and physiological properties of the lizard jaw system, a detailed analysis of the structure of the jaw apparatus was undertaken in the insectivorous lizard Plocederma stellio and in closely related herbivorous lizards of the genus Uromastix. The morphological and physiological properties of the jaw system in P. stellio and U. aegyptius were studied by means of dissections, light microscopy, histochemical characterisations, and in vivo stimulation experiments. The skull of Uromastix seems to be built for forceful biting (high, short snout). Additionally, the pterygoid muscle is modified in P. stellio, resulting in an additional force component during static biting. Stimulation experiments indicate that jaw muscles in both species are fast, which is supported by histochemical stainings. However, the oxidative capacity of the jaw muscles is larger in Uromastix. Contraction characteristics and performance of the feeding system (force output) are clearly thermally dependent. We conclude that several characteristics of the jaw system (presence of extra portion of the pterygoid muscle, large oxidative capacity of jaw muscles) in Uromastix may be attributed to its herbivorous diet. Jaw muscles, however, are still faster than expected. This is presumably the result of trade-offs between the thermal characteristics of the jaw adductors and the herbivorous lifestyle of these animals.

Animals↗

Cranial kinesis in gekkonid lizards

Cranial kinesis was studied in two species of gekkonid lizard, Gekko gecko and Phelsuma madagascariensis, using cineradiography and electromyography. The skull of these geckoes showed the three types of kinesis described by Versluys at the beginning of this century: streptostyly, mesokinesis and metakinesis. In accordance with the later model of Frazzetta, the skull of these animals can be modelled by a quadratic crank system: when the mouth opens during feeding, the quadrate rotates forward, the palato-maxillary unit is lifted and the occipital unit swings forward. During jaw closing, the inverse movements are observed; during crushing, the system is retracted beyond its resting position. The data gathered here indicate that the coupled kinesis (streptostyly + mesokinesis) is most prominently present during the capture and crushing cycles of feeding and is largely absent during late intraoral transport, swallowing, drinking and breathing. The electromyographic data indicate a consistent pattern of muscular activation, with the jaw opener and pterygoid protractor always active during the fast opening phase, and the jaw closers active during closing and crushing. Our data generally support the model of Frazzetta. Although the data gathered here do not allow speculation on the functional significance of the kinesis, they clearly provide some key elements required for a further investigation of the functional and adaptive basis of the system.

Journal Article↗

Kinematics of intraoral transport and swallowing in the herbivorous lizard uromastix acanthinurus

The kinematics of intraoral transport and swallowing in lizards of the species Uromastix acanthinurus (Chamaeleonidae, Leiolepidinae) were investigated using cineradiography (50 frames s-1). Additional recordings were also made using high-speed (500 frames s-1) and conventional video systems (25 frames s-1). Small metal markers were inserted into different parts of the upper and lower jaw and the tongue. Cineradiographic images were digitised, and displacements of the body, head, upper and lower jaw and the tongue were quantified. Twenty additional variables depicting displacements and the timing of events were calculated. Multivariate analyses of variance indicated significant differences between feeding stages. Remarkably, only very few food-type-dependent differences were observed during intraoral transport, and no such differences could be demonstrated during swallowing. Using previously published data for the closely related insectivorous lizard Plocederma stellio, the effect of dietary specialisation in U. acanthinurus on the kinematic variables while eating locusts was examined. Species differed in a number of gape- and tongue-related variables. These differences may be related to differences in tongue structure between the species. Clearly, U. acanthinurus possesses a specialised gut and dental structure that allows them efficiently to cut pieces from whole leaves. However, a decrease in modulatory capacity seems to be a consequence of dietary specialisation in Uromastix acanthinurus.

Journal Article↗

Tongue flicking in agamid lizards: morphology, kinematics, and muscle activity patterns.

We wanted to examine whether a relation between foraging strategy, morphology, the mechanics of tongue protrusion, and prey chemical detection and discrimination exists in agamid lizards. Tongue-flick behavior was observed in two species of this family: Uromastix acanthinurus and Plocederma stellio. Potential prey chemical discrimination by means of tongue flicking was examined by using applicator tests. Tongue flicks were subsequently recorded by high-speed video in combination with the electrical activity of a number of jaw and hyolingual muscles. The kinematics of jaws and tongue and the muscle activity patterns were quantified. To investigate if the observed differences in tongue-flick behavior (mainly in the frequency of use) are translated into corresponding differences in tongue morphology, the tongues of both species were examined by light and scanning electron microscopy. The species differed mainly in the surface morphology of the foretongue and in the abundance and distribution of taste buds on the tongue and oral cavity. These differences can be related to behavioural observations; whereas U. acanthinurus readily uses tongue flicks to detect and discriminate between food items, P. stellio does not. However, differences in tongue-flick mechanics (kinematics, electromyograms) between both species were minor. Based on the data gathered in this study and from previously published data, an evolutionary transformation series leading to the complex tongue-flick cycles as observed in snakes is proposed. The required morphological and mechanical changes that accompany such an evolutionary sequence are discussed.

Animals↗

Quantitative analysis of jaw and hyolingual muscle activity during feeding in the lizard Agama stellio

The activity of jaw and hyolingual muscles during the entire feeding sequence is examined in the lizard Agama stellio, with special focus on the intraoral transport and swallowing stages. Correlation of electromyography (EMG) data with kinematics shows that the kinematic phases (slow opening, SO; fast opening, FO; fast closing, FC; slow closing/power stroke, SC/PS) are characterised by distinct activities in the jaw and hyolingual muscles. The SO phase is clearly the result of tongue protraction (upon protraction, the tongue is pulled against the prey and consequently the lower jaw is pushed down), whereas the FO phase is caused by activity in the jaw opener and dorsal cervical muscles. Both the FC and SC/PS phases are characterised by pronounced activity in the jaw adductor muscles. Tongue retraction is produced by activity in the hyoid and tongue retractor muscles. A quantitative analysis of time-related EMG data shows that, in accordance with the kinematic analyses, three different stages can be recognised as components of the feeding cycle: prey capture, intraoral transport and swallowing. However, analysis of intensity-related data allowed a fourth stage, crushing, to be detected. Whereas there are indications that prey capture, intraoral transport and swallowing are controlled by different motor patterns, the differences between crushing and transport are likely to be caused by feedback mechanisms. Our results show the importance of including intensity-related data in quantitative analyses of EMG recordings in order to discriminate between feeding stages. Additionally, it is shown that both the jaw and the hyolingual muscles play crucial roles during feeding. During all stages, movements of the hyolingual apparatus are an essential part of the feeding cycle. Thus, when examining lizard feeding mechanisms, the activity patterns of the hyolingual muscles should not be neglected.

Journal Article↗

Kinematics of feeding in the lizard Agama stellio

The kinematics of prey capture, intraoral transport and swallowing in lizards of the species Agama stellio (Agamidae) were investigated using cineradiography (50 frames s-1) and high-speed video recordings (500 frames s-1). Small metal markers were inserted into different parts of the upper and lower jaw and the tongue. Video and cineradiographic images were digitized, and displacements of the body, head, upper and lower jaw and the tongue were quantified. Twenty additional variables depicting displacements and timing of events were calculated. A factor analysis performed on the kinematic data separates prey capture and swallowing cycles from intraoral transport bites. However, the intraoral transport stage cannot be separated into chewing (reduction) and transport bites. The effect of prey type and size on the feeding kinematics of intraoral transport and swallowing cycles was investigated. During the intraoral transport stage, distinct aspects (e.g. durations, maximal excursions) of the gape and tongue cycle are modulated in response to both the size and type of the prey item. The results for A. stellio generally agree with a previous model, although it is the entire slow opening phase rather than solely the duration of the second part of this phase that is affected by the size of the prey. The intraoral transport cycles in A. stellio show the two synapomorphic characteristics of tetrapods (tongue-based terrestrial intraoral prey transport and the existence of a long preparatory period of prey compression). However, not all five characters of the feeding cycle previously proposed for amniotes are present in A. stellio. One major difference is that in A. stellio the recovery of the hyolingual apparatus does not take place during the slow opening phase but during the slow closing/powerstroke phase.

Journal Article↗