Why the chameleon has spiral-shaped muscle fibres in its tongue

The intralingual accelerator muscle is the primary actuator for the remarkable ballistic tongue projection of the chameleon. At rest, this muscle envelopes the elongated entoglossal process, a cylindrically shaped bone with a tapering distal end. During tongue projection, the accelerator muscle elongates and slides forward along the entoglossal process until the entire muscle extends beyond the distal end of the process. The accelerator muscle fibres are arranged in transverse planes (small deviations are possible), and form (hitherto unexplained) spiral-shaped arcs from the peripheral to the internal boundary.To initiate tongue projection, the muscle fibres probably generate a high intramuscular pressure. The resulting negative pressure gradient (from base to tip) causes the muscle to elongate and to accelerate forward. Effective forward sliding is made possible by a lubricant and a relatively low normal stress exerted on the proximal cylindrical part of the entoglossal process. A relatively high normal stress is, however, probably required for an effective acceleration of muscle tissue over the tapered end of the process. For optimal performance, the fast extension movement should occur without significant (energy absorbing) torsional motion of the tongue. In addition, the tongue extension movement is aided by a close packing of the muscles fibres (required for a high power density) and a uniform strain and work output in every cross-section of the muscle.A quantitative model of the accelerator muscle was developed that predicts internal muscle fibre arrangements based on the functional requirements above and the physical principle of mechanical stability. The curved shapes and orientations of the muscle fibres typically found in the accelerator muscle were accurately predicted by the model. Furthermore, the model predicts that the reduction of the entoglossal radius towards the tip (and thus the internal radius of the muscle) tends to increase the normal stress on the entoglossal bone.     

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

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

Morphology and 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.     

The mechanism of dewlap extension in Anolis carolinensis (Reptilia: Iguanidae) with histological analysis of the hyoid apparatus

Anolis carolinensis has two aggressive displays involving movements of the hyoid apparatus: erection of the throat and extension of the dewlap. Erection of the throat is an enlargement of the gular region and dewlap extension consists of a vertical erection of the gular flap. Cinefluoroscopy and high speed cinematography show that the dewlap is extended in three phases: 1) protraction of the entire hyoid apparatus; 2) forward pivoting movement of the ceratobranchials II; and 3) retraction of the ceratobranchials II and the entire hyoid apparatus. The cartilaginous elements of the hyoid apparatus are variably mineralized. The entoglossal process and the hypohyals are the most calcified elements. The mineralized portion of the hyoid body, to which the other elements articulate, presents a complex pattern. The calcification of entoglossal process and the hypohyals stop just where they are fused with the hyoid body. The hyoid body presents four mineralized masses, two central corresponding to the base of the ceratobranchials II and two lateral being the head of the ossified ceratobranchials I. The lateral masses articulate on the central masses by a synovial joint. Morphologically, the ceratobranchials II form the hyoid body and become separated at the mid length of the synovial articulation of the ceratobranchials I and the hyoid body. The calcified matrix of the ceratobranchials II gradually changes from a large calcified mass (within the hyoid body) to a semicircle, opened ventrally, which permits their bending during dewlap extension. The highly mineralized posterior tip of the entoglossal process and the hyoid body serve as a pivot to pivoting forward movement of the ceratobranchials II producing at the change of the pattern of mineralization. Forward movement of the ceratobranchials II is produced by electrical stimulation of the M. branchio hyoideus. The opposition of the throat skin to the movement of the ceratobranchials II produces the bending of those longest elements. Electrical stimulation of the hyoid muscles confirms the key role of M. branchiohyoideus during dewlap extension. Simultaneous contractions of all the hyoid and extrinsic tongue (retractor and protractor) muscles with the M. branchiohyoideus during dewlap extension may be a possible motor pattern for dewlap extension in Anolis lizards.     

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.     

Convergently evolved muscle architecture enables high-performance ballistic movement in salamanders

Elastically powered ballistic movements, such as tongue projection, are common in nature, likely due to benefits such as increased acceleration and distance of movement, and decreased thermal sensitivity imparted by elastic mechanisms. Within Plethodontidae, both muscle-powered and elastically powered ballistic tongue projection occur. Thus, we examine how elastically powered ballistic tongue projection morphology has evolved from muscle powered projection at the level of the projector muscles (m. subarcualis rectus [SAR]). We find that two main SAR morphologies have evolved within Plethodontidae. The first SAR morphology is conducive to elastically powered ballistic projection. This ballistic SAR morphology has evolved multiple, independent times within Plethodontidae, and results from the correlated evolution of several traits including increased collagen aponeuroses, larger SAR muscles, and the loss of inner myofibers attaching directly to the tongue skeleton. While the independent evolution of ballistic SAR morphology has arrived at a similar anatomical design, other tongue structures such as tongue attachment and skeleton folding type varies among species with a ballistic SAR morphology. The second morphology is conducive to muscle-powered projection and is similar to morphology found in an outgroup, Salamandridae. The SAR of these species have inner myofibers that attach to the tongue skeleton, limiting projection distance, coupled with reduced collagen aponeuroses present in relatively small projector muscles. This SAR morphology has likely been retained from ancestors or may be related to feeding ecology. Overall, a ballistic SAR morphology has evolved repeatedly and independently due to the correlated evolution of several SAR traits, including the loss of inner myofibers, which is likely a defining feature of ballistic projection.     

MUSCLE TRADE‐OFFS IN A POWER‐AMPLIFIED PREY CAPTURE SYSTEM

Should animals operating at great speeds and accelerations use fast or slow muscles? The answer hinges on a fundamental trade‐off: muscles can be maximally fast or forceful, but not both. Direct lever systems offer a straightforward manifestation of this trade‐off, yet the fastest organisms use power amplification, not direct lever action. Power‐amplified systems typically use slow, forceful muscles to preload springs, which then rapidly release elastic potential energy to generate high speeds and accelerations. However, a fast response to a stimulus may necessitate fast spring‐loading. Across 22 mantis shrimp species (Stomatopoda), this study examined how muscle anatomy correlates with spring mechanics and appendage type. We found that muscle force is maximized through physiological cross‐sectional area, but not through sarcomere length. Sit‐and‐wait predators (spearers) had the shortest sarcomere lengths (fastest contractions) and the slowest strike speeds. The species that crush shells (smashers) had the fastest speeds, most forceful springs, and longest sarcomeres. The origin of the smasher clade yielded dazzlingly high accelerations, perhaps due to the release from fast spring‐loading for evasive prey capture. This study offers a new window into the dynamics of force–speed trade‐offs in muscles in the biomechanical, comparative evolutionary framework of power‐amplified systems.     

Fatiguing contractions of tongue protrudor and retractor muscles: influence of systemic hypoxia.

The influence of systemic hypoxia on the endurance performance of tongue protrudor and retractor muscles was examined in anesthetized, ventilated rats. Tongue protrudor (genioglossus) or retractor (hyoglossus and styloglossus) muscles were activated via medial or lateral XII nerve branch stimulation (0.1-ms pulse; 40 Hz; 330-ms trains; 1 train/s). Maximal evoked potentials (M waves) of genioglossus and hyoglossus were monitored with electromyography. Fatigue tests were performed under normoxic and hypoxic (arterial PO(2) = 50 +/- 1 Torr) conditions in separate animals. The fatigue index (FI; %initial force) after 5 min of normoxic stimulation was 85 +/- 6 and 79 +/- 7% for tongue protrudor and retractor muscles, respectively; these values were significantly lower during hypoxia (protrudor FI = 52 +/- 10, retractor FI = 18 +/- 6%; P < 0.05). Protrudor and retractor muscle M-wave amplitude declined over the course of the hypoxic fatigue test but did not change during normoxia (P < 0.05). We conclude that hypoxia attenuates tongue protrudor and retractor muscle endurance performance; potential mechanisms include neuromuscular transmission failure and/or diminished sarcolemmal excitability.     

Consequences of periodic augmented breaths on tongue muscle activities in hypoxic rats.

This study was designed to investigate the influence of hypoxia-evoked augmented breaths (ABs) on respiratory-related tongue protrudor and retractor muscle activities and inspiratory pump muscle output. Genioglossus (GG) and hyoglossus (HG) electromyogram (EMG) activities and respiratory-related tongue movements were compared with peak esophageal pressure (Pes; negative change in pressure during inspiration) and minute Pes (Pes x respiratory frequency = Pes/min) before and after ABs evoked by sustained poikilocapnic, isocapnic, and hypercapnic hypoxia in spontaneously breathing, anesthetized rats. ABs evoked by poikilocapnic and isocapnic hypoxia triggered long-lasting (duration at least 10 respiratory cycles) reductions in GG and HG EMG activities and tongue movements relative to pre-AB levels, but Pes was reduced transiently (duration of <10 respiratory cycles) after ABs. Adding 7% CO(2) to the hypoxic inspirate had no effect on the frequency of evoked ABs, but this prevented long-term declines in tongue muscle activities. Bilateral vagotomy abolished hypoxia-induced ABs and stabilized drive to the tongue muscles during each hypoxic condition. We conclude that, in the rat, hypoxia-evoked ABs 1) elicit long-lasting reductions in protrudor and retractor tongue muscle activities, 2) produce short-term declines in inspiratory pump muscle output, and 3) are mediated by vagal afferents. The more prolonged reductions in pharyngeal airway vs. pump muscle activities may lead to upper airway narrowing or collapse after spontaneous ABs.     

Episodic hypoxia induces long-term facilitation of neural drive to tongue protrudor and retractor muscles.

Hypoxic episodes can evoke a prolonged augmentation of inspiratory motor output called long-term facilitation (LTF). Hypoglossal (XII) LTF has been assumed to represent increased tongue protrudor muscle activation and pharyngeal airway dilation. However, recent studies indicate that tongue protrudor and retractor muscles are coactivated during inspiration, a behavior that promotes upper airway patency by reducing airway compliance. These experiments tested the hypothesis that XII LTF is manifest as increased inspiratory drive to both tongue protrudor and retractor muscles. Neurograms were recorded in the medial XII nerve branch (XIIMED; contains tongue protrudor motor axons), the lateral XII nerve branch (XIILAT; contains tongue retractor motor axons), and the phrenic nerve in anesthetized, vagotomized, paralyzed, ventilated male rats. Strict isocapnia was maintained for 60 min after five 3-min hypoxic episodes (arterial Po(2) = 35 +/- 2 Torr) or sham treatment. Peak inspiratory burst amplitude showed a persistent increase in XIIMED, XIILAT, and phrenic nerves during the hour after episodic hypoxia (P < 0.05 vs. sham). This effect was present regardless of the quantification method (e.g., % baseline vs. percent maximum); however, comparisons of the relative magnitude of LTF between neurograms (e.g., XIIMED vs. XIILAT) varied with the normalization procedure. There was no persistent effect of episodic hypoxia on inspiratory burst frequency (P > 0.05 vs. sham). These data demonstrate that episodic hypoxia induces LTF of inspiratory drive to both tongue protrudor and retractor muscles and underscore the potential contribution of tongue muscle coactivation to regulation of upper airway patency.     

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.     

Muscle directly meets the vast power demands in agile lizards

Level locomotion in small, agile lizards is characterized by intermittent bursts of fast running. These require very large accelerations, often reaching several times g. The power input required to increase kinetic energy is calculated to be as high as 214 W kg(-1) muscle (+/-20 W kg(-1) s.e.; averaged over the complete locomotor cycle) and 952 W kg(-1) muscle (+/-89 W kg(-1) s.e.; instantaneous peak power). In vitro muscle experiments prove that these exceptional power requirements can be met directly by the lizard's muscle fibres alone; there is no need for mechanical power amplifying mechanisms.     

Ultrastructure of Male Sexual Apparatus of Scutellonema brachyurum

Electron micrographs of serial sections show that the male sexual apparatus of Scutellonema brachyurum includes two morphologically identical spicules. Each is composed of a swollen tubular head, crescentic shaft, and leaf-like blade with membranous velum expanded from the central trunk. The spicules are concave and grooved on the ventral side and convex on the dorsal side near the trunk. The trunk is continuous with the shaft and head. Nerve tissue occupies the core of the spicule and includes a dendritic process which gains access to the exterior via a small pore on the lateral side of the spicule tip. Three protractor and two retractor muscles are associated with each spicule. A sensory accessory piece connects with the tip of the gubernaculum and protrudes from the lower side of the opening of the spicular pouch; it protracts and retracts with the muscularized gubernaculum. The gubernaculum varies from bow-shaped in the distal part to boat-shaped in the mid region. A sac exits beneath the accessory piece as a buffer for its movement. A cuticular guiding bar originating from the dorsal wall of the spicular pouch has a tongue. The ventral surface of the tongue is sclerotized to separate the two spicules. It is mobile by muscles of the protractor gubernaculi, retractor gubernaculi, and seductor gubernaculi.     

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.     

MORPHOLOGY OF THE TONGUE APPARATUS OF C1R1DOPS ANNA (DREPANIDIDAE)

Ciridops possesses a tubular tongue with a fringed tip and lateral sides. Closure of the tube is by overlap of the lateral fringes.
The tongue skeleton possesses an elongated basihyale, parallel paraglossalia with broad, rounded posterior processes and a slight concavity on the dorsolateral surface of the ceratobranchiale.
All the tongue muscles could be described, although some had been damaged. The M. hypoglossus anterior is absent. The M. ceratoglossus and the M. hypoglossus obliquus are large, the latter muscle inserts completely on the basihyale. A newly discovered muscle, the M. thyreohyoideus superior, is described.
The glottal muscles are described, and their actions in opening and closing the glottis are outlined.
Comparison of Ciridops (Drepanidinae) and Loxops (Psittirostrinae) suggests that the Drepanididae are monophyletic. The closest resemblance in morphology of the tongue apparatus is with the cardueline finches, not with the "coerebids".     

Influence of tongue muscle contraction and dynamic airway pressure on velopharyngeal volume in the rat.

The mammalian pharynx is a collapsible tube that narrows during inspiration as transmural pressure becomes negative. The velopharynx (VP), which lies posterior to the soft palate, is considered to be one of the most collapsible pharyngeal regions. I tested the hypothesis that negative transmural pressure would narrow the VP, and that electrical stimulation of extrinsic tongue muscles would reverse this effect. Pressure (-6, -3, 3, and 6 cmH2O) was applied to the isolated pharyngeal airway of anesthetized rats that were positioned in a 4.7-T MRI scanner. The volume of eight axial slices encompassing the length of the VP was computed at each level of pressure, with and without bilateral hypoglossal nerve stimulation (0.1-ms pulse, one-third maximum force, 80 Hz). Negative pressure narrowed the VP, and either whole hypoglossal nerve stimulation (coactivation of protrudor and retractor muscles) or medial nerve branch stimulation (independent activation of tongue protrudor muscles) reversed this effect, with the greatest impact in the caudal one-third of the VP. The dilating effects of medial branch stimulation were slightly larger than whole nerve stimulation. Positive pressure dilated the VP, but tongue muscle contraction did not cause further dilation under these conditions. I conclude that the narrowest and most collapsible segment of the rat pharynx is in the caudal VP, posterior to the tip of the soft palate. Either coactivation of protrudor and retractor muscles or independent contraction of protrudor muscles caused dilation of this region, but the latter was slightly more effective.     

Tongue evolution in the lungless salamanders,family plethodontidae. II. Function and evolutionary diversity

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.     

The mechanism of shell elevation in Haliotis (Mollusca: Gastropoda) and a consideration of the evolution of the hydrostatic skeleton in Mollusca

Early in molluscan evolution, the development of a conical shell with shell or pedal retractor muscles led to the need of a mechanism for the extension of the foot or the raising of the shell. The forces generated during pedal retraction and extension have been studied in Haliotis midae , an easily obtainable and conveniently large archaeogastropod. In the mantle cavity, cephalopedal venous sinus and ventricle pressure pulses were observed during pedal retraction elicited by the shadow withdrawal reflex, but were never present during extension. However, pressure pulses were recorded in the proximal region of the columellar (or shell) muscle, both during retraction and pedal extension. Sections of this region of the muscle show a three dimensional network of muscle fibres, consisting of retractor fibres passing down to the foot and circumferential and radial fibres. Contraction of the two latter sets of fibres would bring about extension of the retractors, without the use of a discrete hydrostatic skeleton, and appears to be the principal mechanism of pedal extension. Similar muscular structures, here termed the muscular antagonistic system, have been observed in the columellar muscle of other gastropods and in the cephalopod mantle. In contrast, this system has not been observed in the proximal region of the pedal retractors of bivalves or scaphopods, for the pedal haemocoel, which allows muscular antagonism in the manner of a classical hydrostatic skeleton, has developed in association with the burrowing habit. The significance of the muscular antagonistic system in molluscan evolution is discussed.     

Evidence for a vertebrate catapult: elastic energy storage in the plantaris tendon during frog jumping

Anuran jumping is one of the most powerful accelerations in vertebrate locomotion. Several species are hypothesized to use a catapult-like mechanism to store and rapidly release elastic energy, producing power outputs far beyond the capability of muscle. Most evidence for this mechanism comes from measurements of whole-body power output; the decoupling of joint motion and muscle shortening expected in a catapult-like mechanism has not been demonstrated. We used high-speed marker-based biplanar X-ray cinefluoroscopy to quantify plantaris muscle fascicle strain and ankle joint motion in frogs in order to test for two hallmarks of a catapult mechanism: (i) shortening of fascicles prior to joint movement (during tendon stretch), and (ii) rapid joint movement during the jump without rapid muscle-shortening (during tendon recoil). During all jumps, muscle fascicles shortened by an average of 7.8 per cent (54% of total strain) prior to joint movement, stretching the tendon. The subsequent period of initial joint movement and high joint angular acceleration occurred with minimal muscle fascicle length change, consistent with the recoil of the elastic tendon. These data support the plantaris longus tendon as a site of elastic energy storage during frog jumping, and demonstrate that catapult mechanisms may be employed even in sub-maximal jumps.     

Combined kinematic and electromyographic analyses of proleg function during crawling by the caterpillar Manduca sexta

The planta retractor muscles in the prolegs of Manduca sexta caterpillars are a frequently-used model system for investigating a number of problems in neurobiology. We have combined kinematic and electromyogram analysis of proleg movements during crawling to examine the roles of these muscles during normal behavior. We found that retractor muscle activity is highly stereotyped, and that the primary function of these muscles is to disengage the crochets at the tip of the proleg for the swing phase of crawling. The duration of activity of the muscles was tightly coupled to the phasing of crawling behavior. The stepping patterns of animals changed to accommodate variations in the substrate, but the relative timing of retractor muscle activity was unaffected. There were no clear correlations between the various properties of motoneuronal input to the muscle (duration of activity, number of spikes, peak frequency of spikes) and the resulting muscle length change. Perhaps because it functions partially as a hydrostat, this may represent a neuromuscular system in which a significant part of the control algorithm is embedded in its morphology.