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1.
Most previous research on metamorphosis of the musculoskeletal system in vertebrates has focused on the transformation of the skeleton. In this paper we focus on the transformation of the muscles of the head during metamorphosis in tiger salamanders ( Ambystoma tigrinum ) in order (1) to provide new data on changes in myology during ontogeny, and (2) to aid in interpreting previous data on the metamorphosis of function in the head of salamanders.
The physiological cross-sectional area of nine head muscles was calculated by measuring fibre angles, fibre lengths, and muscle mass in two samples of tiger salamanders obtained just before and just after metamorphosis. The major mouth-opening muscles (rectus cervicis and depressor mandibulae) exhibit a significant decrease in estimated maximum tetanic tension (MTT) across metamorphosis of about 36%. The jaw-closing muscles (adductor mandibulae internus and externus) and the head-lifting muscles (epaxials) also decrease in MTT but not significantly. The muscles associated with tongue projection during feeding on land (the subarcualis rectus I, geniohyoideus, interhyoideus and intermandibularis) all show a slight increase in MTT at metamorphosis.
Metamorphic transformation of feeding behaviour in Ambystoma tigrinum involves changes in performance, the design of skeletal elements, changes in muscle force-generating capability, and changes in hydrodynamic design from unidirectional flow in larvae to bidirectional flow during aquatic feeding after metamorphosis. Although muscle activity patterns during aquatic feeding do not change across metamorphosis, tongue-based terrestrial feeding involves a suite of novel muscle activity patterns, morphological characters acquired at metamorphosis, and a metamorphic increase in the masses of muscles important in tongue projection. 相似文献
The physiological cross-sectional area of nine head muscles was calculated by measuring fibre angles, fibre lengths, and muscle mass in two samples of tiger salamanders obtained just before and just after metamorphosis. The major mouth-opening muscles (rectus cervicis and depressor mandibulae) exhibit a significant decrease in estimated maximum tetanic tension (MTT) across metamorphosis of about 36%. The jaw-closing muscles (adductor mandibulae internus and externus) and the head-lifting muscles (epaxials) also decrease in MTT but not significantly. The muscles associated with tongue projection during feeding on land (the subarcualis rectus I, geniohyoideus, interhyoideus and intermandibularis) all show a slight increase in MTT at metamorphosis.
Metamorphic transformation of feeding behaviour in Ambystoma tigrinum involves changes in performance, the design of skeletal elements, changes in muscle force-generating capability, and changes in hydrodynamic design from unidirectional flow in larvae to bidirectional flow during aquatic feeding after metamorphosis. Although muscle activity patterns during aquatic feeding do not change across metamorphosis, tongue-based terrestrial feeding involves a suite of novel muscle activity patterns, morphological characters acquired at metamorphosis, and a metamorphic increase in the masses of muscles important in tongue projection. 相似文献
2.
The subarcualis rectus I muscle (SAR) in the feeding mechanism of four tiger salamanders (Ambystoma tigrinum) was removed early in ontogeny and these individuals were allowed to complete metamorphosis. This procedure resulted in postmetamorphic tiger salamanders which differed from control individuals in the size (and thus force generating capacity) of the SAR muscle. The experimental manipulation of muscle ontogeny allowed a test of previous hypotheses of SAR function in postmetamorphic individuals. Multivariate analysis of variance for kinematic variables measured from high-speed video records of feeding revealed that experimentally modified tiger salamanders did not protract the hyobranchial apparatus or project the tongue from the mouth during feeding. Removal of the SAR muscle resulted in significantly reduced hyobranchial elevation in the buccal cavity and reduced maximum tongue projection distance. 相似文献
3.
This study provides the first data on muscle activity patterns during active feeding in a larval anuran. Data regarding muscle function during gill irrigation and hyperexpiration are also provided. Electromyographic and kinematic data were recorded from six mandibular and hyoid muscles in unanesthetized, unrestrained larvae of Rana catesbeiana. Only three (hyoangularis, orbitohyoideus, anterior interhyoideus) of the six muscles examined are active during gill irrigation. Feeding cycles are characterized by the recruitment of three additional muscles: intermandibularis, suspensorioangularis, and levator mandibulae longus superficialis. The latter two contribute, respectively, to wide opening and forceful closing of the mouth during feeding. Hyperexpiration is characterized by a reversal of water flow anteriorly out of the mouth. This hydrodynamic change occurs due to modulation of the timing of firing of the anterior interhyoideus, as well as recruitment of the posterior interhyoideus, which is only active during hyperexpiration. Both regions of the interhyoideus, which are responsible for evacuation of the buccal cavity, are active during the opening phase of hyperexpiration. Kinematically, transitioning from gill irrigation to feeding involves both an overall shortening of the gape cycle and a shift in the relative length of opening phase vs. closing phase. Our results corroborate many of the findings of Gradwell ([1972] Can J Zool 50:501-521) regarding muscle function during gill irrigation and hyperexpiration. Furthermore, we demonstrate that in larval anurans the transition from gill irrigation to feeding involves modulation of gape cycle kinematics, changes in the level of activity of muscles, and recruitment of muscles that are not active during irrigation. In light of new data presented here, a review of muscle function in tadpoles is also provided. 相似文献
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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. 相似文献
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The Hynobiidae is generally regarded as the most phylogenetically basal and least derived extant family of terrestrial salamanders. As in the other families of terrestrial salamanders, prey capture in the Hynobiidae is accomplished by lingual prehension. In Hynobius, the prey capture system appears to be a mosaic of derived and primitive features. This, in conjunction with previous studies, suggests that the hyolingual systems of all families of terrestrial salamanders have evolved various degrees of specialization since the appearance of the common ancestral condition. We propose that the generalized feeding system for the extant terrestrial salamanders includes a hyolingual skeleton comprised of one basibranchial, one pair of radial or radial-like structures, two pairs of ceratobranchials, two pairs of epibranchials, one pair of ceratohyals, and one urohyal arranged in a configuration similar to that of Hynobius; a simple, sac-like secondary tongue pad; a lift and thrust system of tongue projection; a four-part gape cycle; and a forward head and body surge. Modifications to this general plan, previously described for the disparate families, include various changes in the size, shape, and definition of the tongue pad, changes in the specific types of structures and configurations in the anterior hyolingual skeleton, secondary ossification in the posterior hyolingual skeleton, the appearance of various protrusion, projection, and flipping systems for tongue protraction, simplification of the kinematic gape profile, and loss of the forward head and body surge. The evolutionary trends in these modifications have provided a rich data set from which much phylogenetic information has been inferred. © 1996 Wiley-Liss, Inc. 相似文献
8.
Stephen M. Reilly George V. Lauder 《Evolution; international journal of organic evolution》1990,44(6):1542-1557
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. 相似文献
9.
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. 相似文献
10.
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. 相似文献
11.
Amphibians (Lissamphibia) are characterized by a bi‐phasic life‐cycle that comprises an aquatic larval stage and metamorphosis to the adult. The ancestral aquatic feeding behavior of amphibian larvae is suction feeding. The negative pressure that is needed for ingestion of prey is created by depression of the hyobranchial apparatus as a result of hyobranchial muscle action. Understanding the homologies of hyobranchial muscles in amphibian larvae is a crucial step in understanding the evolution of this important character complex. However, the literature mostly focuses on the adult musculature and terms used for hyal and ventral branchial muscles in different amphibians often do not reflect homologies across lissamphibian orders. Here we describe the hyal and ventral branchial musculature in larvae of caecilians (Gymnophiona) and salamanders (Caudata), including juveniles of two permanently aquatic salamander species. Based on previous alternative terminology schemes, we propose a terminology for the hyal and ventral branchial muscles that reflects the homologies of muscles and that is suited for studies on hyobranchial muscle evolution in amphibians. We present a discussion of the hyal and ventral branchial muscles in larvae of the most recent common ancestor of amphibians (i.e. the ground plan of Lissamphibia). Based on our terminology, the hyal and ventral branchial musculature of caecilians and salamanders comprises the following muscles: m. depressor mandibulae, m. depressor mandibulae posterior, m. hyomandibularis, m. branchiohyoideus externus, m. interhyoideus, m. interhyoideus posterior, m. subarcualis rectus I, m. subarcualis obliquus II, m. subarcualis obliquus III, m. subarcualis rectus II‐IV, and m. transversus ventralis IV. Except for the m. branchiohyoideus externus, all muscles considered herein can be assigned to the ground plan of the Lissamphibia with certainty. The m. branchiohyoideus externus is either apomorphic for the Batrachia (frogs + salamanders) or salamander larvae depending on whether or not a homologous muscle is present in frog tadpoles. J. Morphol., 2011. © 2011 Wiley‐Liss, Inc. 相似文献
12.
While ontogenetic analyses of skull development have contributed to our understanding of phylogenetic patterns in vertebrates, there are few studies of taxa that undergo a relatively discrete and rapid change in morphology during development (metamorphosis). Morphological changes occurring in the head at metamorphosis in tiger salamanders (Ambystoma tigrinum) were quantified by a morphometric analysis of cranial osteology and myology to document patterns of change during metamorphosis. We employed a cross-sectional analysis using a sample of larvae just prior to metamorphosis and a sample of transformed individuals just after metamorphosis, as well as larvae undergoing metamorphosis. There were no differences in external size of the head among the larval and transformed samples. The hyobranchial apparatus showed many dramatic changes at metamorphosis, including shortening of ceratobranchial 1 and the basibranchial. The subarcualis rectus muscle increased greatly in length at metamorphosis, as did hypobranchial length and internasal distance. A truss analysis of dorsal skull shape showed that at metamorphosis the snout becomes wider, the maxillary and squamosal triangles rotate posteromedially, and the neurocranium shortens (while maintaining its width), resulting in an overall decrease in skull length at metamorphosis. These morphometric differences are interpreted in light of recent data on the functional morphology of feeding in salamanders. Morphological reorganization of the hyobranchial apparatus and shape changes in the skull are related to the acquisition of a novel terrestrial feeding mode (tongue projection) at metamorphosis. Metamorphic changes (both internal and external) that can be used to judge metamorphic condition are discussed. 相似文献
13.
Jenna A. Monroy Kiisa C. Nishikawa 《Journal of comparative physiology. A, Neuroethology, sensory, neural, and behavioral physiology》2009,195(9):843-852
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. 相似文献
14.
K. C. NISHIKAWA D. C. CANNATELLA F.L.S. 《Zoological Journal of the Linnean Society》1991,103(3):289-307
The kinematics of prey capture by Ascaphus truei was investigated. High-speed films (100 fps) of 13 successful and one unsuccessful prey capture sequences from six adult frogs were analysed. Ascaphus , the sister group of all living frogs, shares several aspects of feeding kinematics, including rotation of the tongue pad about the mandibular symphysis and mandibular bending during mouth opening and closing, with more derived frogs such as Bufo marinus. The times required for tongue retraction, mouth opening and closing are similar in Ascaphus and Bufo. However, because Bufo is much larger and protracts its tongue much farther than Ascaphus , the velocities of tongue retraction, mouth opening and mouth closing are relatively lower in Ascaphus than in Bufo. Differences in prey capture between Ascaphus and Bufo marinus are (1) the distance of tongue protraction is less in Ascaphus (±0.5 cm) than in Bufo (c. 2 cm); and (2) lunging of the whole body is more pronounced in Ascaphus. Prey capture is highly variable in Ascaphus. An intraoral transport sequence is sometimes (7 of 14 observations) inserted into the prey capture cycle before the completion of mouth closing. The gape cycles range from 80–150 ms for sequences with no oral transport and from 130–280 ms for sequences with oral transport. Also, the time required for tongue retraction is significantly longer in the unsuccessful capture attempt. This variability is generally greater than that observed during prey capture in salamanders, and suggests that frogs and salamanders may differ in the importance of sensory feedback in coordinating prey capture. 相似文献
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16.
A recently presented model of tongue projection dynamics is used to generate a series of predictions concerning morphologies to be expected under selection for increased distance of projection, increased speed of projection, and increased directional versatility. A general understanding of biomechanical events and the model are used as points of departure for making specific predictions concerning details of structure in skeletal, muscular and connective tissue components of the tongue and associated structures. Comparative methods are used to examine these predictions in the genera of plethodontid salamanders. These salamanders are known to project their tongues to different degrees, and this knowledge is used to test the hypotheses concerning morphological specialization. Three distinct groups of plethodontid salamanders have evolved specializations for long distance projection, and these genera differ from one another in important ways in respect to specific character complexes. For example, the tropical genera and Hydromantes use CBII as the major force transmission element in the skeleton, while Eurycea and its allies use CBI in this role. Hydromantes differs from both in having a uniquely proportioned and structured hyobranchial skeleton and associated musculature. Less extreme specializations for tongue projection are found in different combinations in three other groups. Finally, two distinct groups of generalized species having only limited tongue projection capabilities are recognized, each having a unique complex of inter-related features. Each of these eight groups is recognized and characterized as a functional mode, and hypotheses concerning the biomechanical meaning of the character complexes of each are formulated. 相似文献
17.
Some species of Clariidae (air breathing catfishes) have extremely large (hypertrophied) jaw closure muscles. Besides producing higher bite forces, the enlarged muscles may also cause higher accelerations of the lower jaw during rapid mouth closure. Thus, jaw adductor hypertrophy could potentially also enable faster mouth closure. In this study, a forward dynamic model of jaw closing is developed to evaluate the importance of jaw adductor hypertrophy on the speed of mouth closure. The model includes inertia, pressure, tissue resistance and hydrodynamic drag forces on the lower jaw, which is modelled as a rotating half-ellipse. Simulations are run for four clariid species showing a gradual increase in jaw adductor hypertrophy (Clarias gariepinus, Clariallabes longicauda, Gymnallabes typus and Channallabes apus). The model was validated using data from high-speed videos of prey captures in these species. In general, the kinematic profiles of the fastest mouth closure from each species are reasonably well predicted by the model. The model was also used to compare the four species during standardized mouth closures (same initial gape angle, travel distance and cranial size). These simulations suggest that the species with enlarged jaw adductors have an increased speed of jaw closure (in comparison with the non-hypertrophied C. gariepinus) for short lower jaw rotations and when feeding at high gape angles. Consequently, the jaw system in these species seems well equipped to capture relatively large, evasive prey. For prey captures during which the lower jaw rotates freely over a larger distance before impacting the prey, the higher kinematic efficiency of the C. gariepinus jaw system results in the fastest jaw closures. In all cases, the model predicts that an increase in the physiological cross-sectional area of the jaw muscles does indeed contribute to the speed of jaw closure in clariid fish. 相似文献
18.
A. Herrel M. Verstappen F. De Vree 《Journal of comparative physiology. A, Neuroethology, sensory, neural, and behavioral physiology》1999,184(5):501-518
The kinematics of jaws and tongue, and jaw muscle activity patterns were investigated in the omnivorous lizard Tiliqua rugosa, and the herbivorous Corucia zebrata (Scincidae) during feeding. Small metal markers were inserted into different parts of the skull, the jaws, and the tongue.
Video and cineradiographic images were digitized and displacements of the head, jaws, and tongue were quantified. Additionally,
muscle activity patterns were recorded, digitized and several variables were determined quantitatively. The effect of food
type on the jaw and hyolingual movement patterns and the jaw muscle activity patterns was investigated for both species. The
kinematic data indicate that distinct aspects of gape and tongue cycles are modulated in response to the food characteristics.
Similarly, in both species, muscle activity patterns are altered in response to the type of food eaten. A comparison of kinematic
and electromyographic patterns during intraoral transport cycles for both species shows that these can be related to food
characteristics such as toughness and mobility. Differences between both species in the response to changes in food characteristics
are minor. Clearly both species are able to fine tune the activation of the jaw muscles, resulting in the appropriate movement
patterns for the type of food eaten.
Accepted: 30 January 1999 相似文献
19.
H. Bradley Shaffer George V. Lauder 《Evolution; international journal of organic evolution》1985,39(1):83-92
Patterns of variation in the feeding mechanism of three species of ambystomatid salamanders (Ambystoma dumerilii, A. mexicanum, and A. ordinarium) were studied to provide insight into the nature of variation in kinematic parameters of the jaw mechanism associated with prey capture. A nested analysis of variance design provided an assessment of the amount of variation in six kinematic variables (measured from 200 frames/sec films of feeding behavior) both among species and among individuals within species. For all six variables, a highly significant proportion of the variance was explained at the intraspecific level. Among species, the most robust discriminators were variables associated with movement of the hyoid. The variables reflecting gape and lifting of the head provided no significant discrimination among species and had large error variances. The hyoid apparatus is the most phylogenetically conservative component of the feeding mechanism in lower vertebrates and was the most stereotyped component of feeding behavior within the salamander species studied here. 相似文献
20.
Functional and evolutionary morphology of lingual feeding in squamate reptiles: phylogenetics and kinematics 总被引:3,自引:0,他引:3
Use of the tongue as a prehensile organ during the ingestion stage of feeding in lizards was studied cinegraphically in seven species. Within Squamata, lingual prehension is limited to a single clade, the Iguania (Iguanidae, Agamidae and Chamaeleontidae), which includes all 'fleshy-tongued' lizards. All remaining squamates (Scleroglossa) use the jaws alone for prey prehension. Lingual prehension and a 'fleshy' tongue are primitive squamate characteristics. Kinematically, lingual ingestion cycles are similar to previously described transport cycles in having slow open, fast open, fast close and slow close-power stroke phases. Tongue movements are sequentially correlated with jaw movements as they are in transport. However, during ingestion, anterior movement of the tongue includes an extra-oral, as well as intra-oral component. Tongue protrusion results in a pronounced slow open-II phase at a large gape distance. A high degree of variability in quantitative aspects of ingestion and transport cycles suggests that modulation through sensory feedback is an important aspect of lizard feeding. Preliminary evidence indicates an important role for hyoid movement in tongue protrusion. Our results are consistent with the Bramble & Wake (1985) model generalized feeding cycle and support their contention that specialized feeding mechanisms often represent modifications of a basic pattern, particularly modification of the slow open phase. 相似文献