The limbs of running mammals are thought to function as inverted struts. When mammals run at constant speed, the ground reaction force vector appears to be directed near the point of rotation of the limb on the body such that there is little or no moment at the joint. If this is true, little or no external work is done at the proximal joints during constant-speed running. This possibility has important implications to the energetics of running and to the coupling of lung ventilation to the locomotor cycle. To test if the forelimb functions as an inverted strut at the shoulder during constant-speed running and to characterize the locomotor function of extrinsic muscles of the forelimb, we monitored changes in the recruitment of six muscles that span the shoulder (the m. pectoralis superficialis descendens, m. pectoralis profundus, m. latissimus dorsi, m. omotransversarius, m. cleidobrachialis and m. trapezius) to controlled manipulations of locomotor forces and moments in trotting dogs (Canis lupus familiaris Linnaeus 1753). Muscle activity was monitored while the dogs trotted at moderate speed (approximately 2 m s(-1)) on a motorized treadmill. Locomotor forces were modified by (1) adding mass to the trunk, (2) inclining the treadmill so that the dogs ran up- and downhill (3) adding mass to the wrists or (4) applying horizontally directed force to the trunk through a leash. When the dogs trotted at constant speed on a level treadmill, the primary protractor muscles of the forelimb exhibited activity during the last part of the ipsilateral support phase and the beginning of swing phase, a pattern that is consistent with the initiation of swing phase but not with active protraction of the limb during the beginning of support phase. Results of the force manipulations were also consistent with the protractor muscles initiating swing phase and contributing to active braking via production of a protractor moment on the forelimb when the dogs decelerate. A similar situation appears to be true for the major retractor muscles of the forelimb. The m. pectoralis profundus and the m. latissimus dorsi were completely silent during the support phase of the ipsilateral limb when the dogs ran unencumbered and exhibited little or no increase in activity when the dogs carried added mass on their backs to increase any retraction torque during the support phase of constant-speed running. The most likely explanation for these observations is that the ground force reaction vector is oriented very close to the fulcrum of the forelimb such that the forelimb functions as a compliant strut at the shoulder when dogs trot at constant speed on level surfaces. Because the moments at the fulcrum of the pectoral girdle appear to be small during the support phase of a trotting step, a case can be made that it is the activity of the extrinsic appendicular muscles that produce the swing phase of the forelimb that explain the coupled phase relationship between ventilatory airflow and the locomotor cycle in trotting dogs.

Many plethodontid salamanders project their tongues ballistically at high speed and for relatively great distances. Capturing evasive prey relies on the tongue reaching the target in minimum time, therefore it is expected that power production, or the rate of energy release, is maximized during tongue launch. We examined the dynamics of tongue projection in three genera of plethodontids (Bolitoglossa, Hydromantes and Eurycea), representing three independent evolutionary transitions to ballistic tongue projection, by using a combination of high speed imaging, kinematic and inverse dynamics analyses and electromyographic recordings from the tongue projector muscle. All three taxa require high-power output of the paired tongue projector muscles to produce the observed kinematics. Required power output peaks in Bolitoglossa at values that exceed the greatest maximum instantaneous power output of vertebrate muscle that has been reported by more than an order of magnitude. The high-power requirements are likely produced through the elastic storage and recovery of muscular kinetic energy. Tongue projector muscle activity precedes the departure of the tongue from the mouth by an average of 117 ms in Bolitoglossa, sufficient time to load the collagenous aponeuroses within the projector muscle with potential energy that is subsequently released at a faster rate during tongue launch.

Biotic factors such as body size and shape have long been known to influence kinematics in vertebrates. Movement in aquatic organisms can also be strongly affected by abiotic factors such as the viscosity of the medium. We examined the effects of both biotic factors and abiotic factors on buccal pumping kinematics in Xenopus tadpoles using high-speed imaging of an ontogenetic series of tadpoles combined with experimental manipulation of the medium over a 10-fold range of viscosity. We found influences of both biotic and abiotic factors on tadpole movements; absolute velocities and excursions of the jaws and hyoid were greater in higher viscosity fluid but durations of movements were unaffected. Smaller tadpoles have relatively wider heads and more robust hyoid muscles used in buccal expansion and compression. Lever arm ratios were found to be constant at all sizes; therefore, smaller tadpoles have relatively higher resolved muscle forces and, like tadpoles in more viscous medium, displayed higher absolute velocities of jaw and hyoid movements. Nonetheless, small tadpoles drew in water at lower Reynolds numbers (Re) than predicted by kinematics, due to negative allometry of the buccal pump. Finally, tadpoles transitioned from a flow regime dominated by viscous forces (Re=2) to an intermediate regime (Re=106).

Environmental temperature impacts the physical activity and ecology of ectothermic animals through its effects onmuscle contractile physiology. Sprinting, swimming, and jumping performance of ecto-therms decreases by at least 33% over a 10 degrees C drop, accompanied by a similar decline inmuscle power. We propose that ballistic movements that are powered by recoil of elastic tissues are less thermally dependent than movements that rely on direct muscular power. We found that an elastically powered movement, ballistic tongue projection in chameleons, maintains high performance over a 20 degrees C range. Peak velocity and power decline by only 10%-19% with a 10 degrees C drop, compared to > 42% for nonelastic, muscle-powered tongue retraction. These results indicate that the elastic recoil mechanism circumvents the constraints that low temperature imposes on muscle rate properties and thereby reduces the thermal dependence of tongue projection. We propose that organisms that use elastic recoil mechanisms for ecologically important movements such as feeding and locomotion may benefit from an expanded thermal niche.

The activity of seven trunk muscles was recorded at two sites along the trunk in adult spotted salamander, Ambystoma maculatum, during swimming and during trotting in water and on land. Several muscles showed patterns of activation that are consistent with the muscles producing a traveling wave of lateral bending during swimming and a standing wave of bending during aquatic and terrestrial trotting: the dorsalis trunci, subvertebralis lateralis and medialis, rectus lateralis and obliquus internus. The interspinalis showed a divergent pattern and was active out of phase with the other muscles suggesting that it functions in vertebral stabilization rather than lateral bending. The obliquus internus and rectus abdominis showed bilateral activity indicating that they counteract sagittal extension of the trunk that is produced when the large dorsal muscles are active to produce lateral bending. Of the muscles examined, only the obliquus internus showed a clear shift in function from lateral bending during swimming to resistance of long-axis torsion during trotting. During terrestrial trotting, muscle recruitment was greater in several muscles than during aquatic trotting, despite similar temporal patterns of muscle activation, suggesting that the trunk is stiffened during terrestrial locomotion against greater gravitational forces whereas the basic functions of the trunk muscles in trotting are conserved across environments.

Extremely specialized and long tongues used for prey capture have evolved independently in plethodontid salamanders and chameleons. In both systems, the demands on tongue projection are probably similar: to maximize projection velocity and distance. Consequently, many of the design features of the projection system in these two groups have converged to an astonishing degree. Both involve the use of power amplification systems based on helically wound muscle fibers that load internal connective tissue sheets as illustrated in previous studies. Demands imposed on tongue retraction, however, are different to some degree. Although in both groups there is a clear demand for retraction capacity ( given the long projection distances), in chameleons there is an added demand for force because they eat large and heavy prey. as indicated by our data, plethodontid salamanders have extremely long tongue retractors with normal striated muscle. Chameleons, on the other hand, evolved long retractors of the supercontracting type. Interestingly, our data show that at least in chameleons, the extreme design of the tongue in function of prey capture appears to have consequences on prey transport, resulting in an increased dependence on the hyoid. In turn, this has lead to an increase in transport-cycle duration and an increase in the number of cycles needed to transport prey in comparison with closely related agamid lizards. Clearly, extreme morphological specializations are tuned to functional and ecological demands and may induce a reduced performance in other functions performed by the same set of integrated structures.