A comparison of the moment arms of pelvic limb muscles in horses bred for acceleration (Quarter Horse) and endurance (Arab)

Selective breeding for performance has resulted in distinct breeds of horse, such as the Quarter Horse (bred for acceleration) and the Arab (bred for endurance). Rapid acceleration, seen during Quarter Horse racing, requires fast powerful muscular contraction and the generation of large joint torques, particularly by the hind limb muscles. This study compared hind limb moment arm lengths in the Quarter Horse and Arab. We hypothesized that Quarter Horse hind limb extensor muscles would have longer moment arms when compared to the Arab, conferring a greater potential for torque generation at the hip, stifle and tarsus during limb extension. Six Quarter Horse and six Arab hind limbs were dissected to determine muscle moment arm lengths for the following muscles: gluteus medius, biceps femoris, semitendinosus, vastus lateralis, gastrocnemius (medialis and lateralis) and tibialis cranialis. The moment arms of biceps femoris (acting at the hip) and gastrocnemius lateralis (acting at the stifle) were significantly longer in the Quarter Horse, although the length of the remaining muscle moment arms were similar in both breeds of horse. All the Quarter Horse muscles were capable of generating greater muscle moments owing to their greater physiological cross-sectional area (PCSA) and therefore greater isometric force potential, which suggests that PCSA is a better determinant of muscle torque than moment arm length in these two breeds of horse. With the exception of gastrocnemius and tibialis cranialis, the observed muscle fascicle length to moment arm ratio (MFL : MA ratio) was greater for the Arab horse muscles. It appears that the Arab muscles have the potential to operate at slower velocities of contraction and hence generate greater force outputs when compared to the Quarter Horse muscles working over a similar range of joint motion; this would indicate that Arab hind limb muscles are optimized to function at maximum economy rather than maximum power output.     

Functional specialisation of pelvic limb anatomy in horses (Equus caballus)

We provide quantitative anatomical data on the muscle-tendon units of the equine pelvic limb. Specifically, we recorded muscle mass, fascicle length, pennation angle, tendon mass and tendon rest length. Physiological cross sectional area was then determined and maximum isometric force estimated. There was proximal-to-distal reduction in muscle volume and fascicle length. Proximal limb tendons were few and, where present, were relatively short. By contrast, distal limb tendons were numerous and long in comparison to mean muscle fascicle length, increasing potential for elastic energy storage. When compared with published data on thoracic limb muscles, proximal pelvic limb muscles were larger in volume and had shorter fascicles. Distal limb muscle architecture was similar in thoracic and pelvic limbs with the exception of flexor digitorum lateralis (lateral head of the deep digital flexor), the architecture of which was similar to that of the pelvic and thoracic limb superficial digital flexors, suggesting a functional similarity.     

Functional specialisation of the pelvic limb of the hare (Lepus europeus)

We provide quantitative anatomical data on the muscle-tendon architecture of the hare pelvic limb (specifically muscle mass, fascicle length, pennation angle, tendon mass and length). In addition, moment arms of major pelvic limb muscles were measured. Maximum isometric force and power of muscles, the moment of force about a joint, and tendon stress and strain were estimated. Data are compared with published data for other cursorial mammals such as the horse and dog, and a non-specialised Lagamorph, the rabbit. The pelvic limb of the hare was found to contain substantial amounts of hip extensor and adductor/abductor muscle volume, which is likely to be required for power production and stability during rapid turning. A proximal to distal reduction in muscle volume and fascicle length was also observed, as is the case in other cursorial quadrupeds, along with a reduction in distal limb mass via the replacement of muscle volume by long distal limb tendons, capable of storing large amounts of elastic energy. The majority of hare pelvic limb muscle moment arms varied with joint position, giving the hare the capacity to vary muscle function with limb posture and presumably different locomotor activities.     

Functional anatomy and muscle moment arms of the pelvic limb of an elite sprinting athlete: the racing greyhound (Canis familiaris)

We provide quantitative anatomical data on the muscle-tendon architecture and geometry of the pelvic limb of an elite sprint athlete, the racing greyhound. Specifically, muscle masses, muscle lengths, fascicle lengths, pennation angles and muscle moment arms were measured. Maximum isometric force and power of muscles, the maximum muscle torque at joints and tendon stress and strain were estimated. We compare data with that published for a generalized breed of canid, and other cursorial mammals such as the horse and hare. The pelvic limb of the racing greyhound had a relatively large volume of hip extensor muscle, which is likely to be required for power production. Per unit body mass, some pelvic limb muscles were relatively larger than those in less specialized canines, and many hip extensor muscles had longer fascicle lengths. It was estimated that substantial extensor moments could be created about the tarsus and hip of the greyhound allowing high power output and potential for rapid acceleration. The racing greyhound hence possesses substantial specializations for enhanced sprint performance.     

Muscle architecture and functional anatomy of the pelvic limb of the ostrich (Struthio camelus)

The functional anatomy of the pelvic limb of the ostrich (Struthio camelus) was investigated in order to assess musculoskeletal specialization related to locomotor performance. The pelvic limbs of ten ostriches were dissected and detailed measurements of all muscle tendon units of the pelvic limb were made, including muscle mass, muscle length, fascicle length, pennation angle, tendon mass and tendon length. From these measurements other muscle properties such as muscle volume, physiological cross-sectional area (PCSA), tendon cross-sectional area, maximum isometric muscle force and tendon stress were derived, using standard relationships and published muscle data. Larger muscles tended to be located more proximally and had longer fascicle lengths and lower pennation angles. This led to an expected proximal to distal reduction in total muscle mass. An exception to this trend was the gastrocnemius muscle, which was found to have the largest volume and PCSA and also had the highest capacity for both force and power production. Generally high-power muscles were located more proximally in the limb, while some small distal muscles (tibialis cranialis and flexor perforatus digiti III), with short fibres, were found to have very high force generation capacities. The greatest proportion of pelvic muscle volume was for the hip extensors, while the highest capacity for force generation was observed in the extensors of the ankle, many of which were also in series with long tendons and thus were functionally suited to elastic energy storage.     

Functional anatomy and muscle moment arms of the thoracic limb of an elite sprinting athlete: the racing greyhound (Canis familiaris)

We provide quantitative muscle–tendon architecture and geometry data for the racing greyhound thoracic limb. Muscle mass, belly length, fascicle lengths, pennation angles and moment arms were measured, as were tendon masses and lengths. Maximum isometric force and maximum power were estimated for muscles, and maximum stress and strain were estimated for tendons. Results are compared with other fast quadrupedal runners, and to previously published data in mixed‐breed dogs. The implications of the functional adaptations of the greyhound thoracic limb for sprinting performance are discussed. The thoracic limb was found to benefit from a similar proportion of locomotor muscle mass to the pelvic limb, suggesting that it may be used to some extent in propulsion, or alternatively that stabilisation is very important in this animal. Extrinsic muscles, especially latissimus dorsi and pectoralis profundus, were predicted to be powerful and important for generating net positive work during accelerations. Proximal biarticular muscles show specialisation toward preventing collapse of the shoulder and elbow joints to enable strut‐like limb function, or some form of dynamic control. Distal muscles did not appear specialised for elastic energy storage, a functional difference to pelvic limb muscles, and the equivalents in horse thoracic limbs. The greyhound thoracic limb appears to possess substantial differences from both that of more ‘sub‐maximal specialist’ quadrupeds, and from the greyhound pelvic limb.     

Variety,sex and ontogenetic differences in the pelvic limb muscle architectural properties of leghorn chickens (Gallus gallus domesticus) and their links with locomotor performance

Leghorn (layer) chickens (Gallus gallus domesticus) differ in locomotor morphology and performance due to artificial selection for standard (large) and bantam (small) varieties, sexual dimorphisms and ontogenetic stage. Here, the hind limb skeletal muscle architectural properties of mature and juvenile standard breeds and mature bantams are compared and linked to measures of locomotor performance. Mature males possessed greater relative muscle physiological cross‐sectional areas (PCSAs) than their conspecific females, indicative of greater force‐generating capacity, and in line with their greater maximum sustainable speeds compared with females. Furthermore, some of the relative fascicle lengths of the pennate muscles were greater in mature males than in mature females, which may permit greater muscle contractibility. Immature standard leghorns, however, did not share the same dimorphisms as their mature forms. The differences in architectural properties between immature and mature standard males indicate that with the onset of male sexual maturity, concomitant with increasing muscle mass in males, the relative fascicle lengths of pennate muscles and the relative PCSAs of the parallel‐fibred muscles also increase. The age‐related differences in standard breed male muscle architecture are linked to the presence and absence of sex differences in maximum aerobic speeds. Males of bantam and standard varieties shared similar muscle proportions (% body mass), but exhibited intrinsic muscle differences with a tendency for greater force‐generating capabilities in bantams and greater contractile capabilities in standards. The metabolic costs associated with the longer fascicle lengths, together with more crouched limbs in standard than in bantam males may explain the lack of allometry in the minimum metabolic cost of transport between these birds of different size.     

Muscle�Ctendon structure and dimensions in adults and children

Muscle performance is closely related to the architecture and dimensions of the muscle–tendon unit and the effect of maturation on these architectural characteristics in humans is currently unknown. This study determined whether there are differences in musculo‐tendinous architecture between adults and children of both sexes. Fascicle length and pennation angle were measured from ultrasound images at three sites along the length of the vastus intermedius, vastus lateralis, vastis medialis and rectus femoris muscles. Muscle volume and muscle–tendon length were measured from magnetic resonance images. Muscle physiological cross‐sectional area (PCSA) was calculated as the ratio of muscle volume to optimum fascicle length. Fascicle length was greater in the adult groups than in children (P < 0.05) but pennation angle did not differ between groups (P > 0.05). The ratios between fascicle and muscle length and between fascicle and tendon length were not different (P > 0.05) between adults and children for any quadriceps muscle. Quadriceps volume and PCSA of each muscle were greater in adults than children (P < 0.01) but the relative proportion of each head to the total quadriceps volume was similar in all groups. However, the difference in PCSA between adults and children (men ～ 104% greater than boys, women ～ 57% greater than girls) was greater (P < 0.05) than the difference in fascicle length (men ～ 37% greater than boys, women ～ 10% greater than girls). It is concluded that the fascicle, muscle and tendon lengthen proportionally during maturation, thus the muscle–tendon stiffness and excursion range are likely to be similar in children and adults but the relatively greater increase in PCSA than fascicle length indicates that adult muscles are better designed for force production than children’s muscles.     

Functional specialization and ontogenetic scaling of limb anatomy in Alligator mississippiensis

Crocodylians exhibit a fascinating diversity of terrestrial gaits and limb motions that remain poorly described and are of great importance to understanding their natural history and evolution. Their musculoskeletal anatomy is pivotal to this diversity and yet only qualitative studies of muscle‐tendon unit anatomy exist. The relative masses and internal architecture (fascicle lengths and physiological cross‐sectional areas) of muscles of the pectoral and pelvic limbs of American alligators (Alligator mississippiensis Daudin 1801) were recorded for an ontogenetic series of wild specimens (n = 15, body masses from 0.5 to 60 kg). The data were analysed by reduced major axis regression to determine scaling relationships with body mass. Physiological cross‐sectional areas and therefore muscle force‐generating capacity were found to be greater in the extensor (anti‐gravity) muscles of the pelvic limb than in the pectoral limb, reflecting how crocodylians differ from mammals in having greater loading of the hindlimbs than the forelimbs. Muscle masses and architecture were generally found to scale isometrically with body mass, suggesting an ontogenetic decrease in terrestrial athleticism. This concurs with the findings of previous studies showing ontogenetic decreases in limb bone length and the general scaling principle of a decline of strength : weight ratios with increasing size in animals. Exceptions to isometric scaling found included positive allometry in fascicle length for extensor musculature of both limbs, suggesting an ontogenetic increase in working range interpreted as increasing postural variability – in particular the major hip extensors – the interpretation of which is complicated by previous described ontogenetic increase of moment arms for these muscles.     

Functional specialisation of the thoracic limb of the hare (Lepus europeus)

We provide quantitative anatomical data on the muscle-tendon architecture of the hare thoracic limb (specifically muscle mass, fascicle length, pennation angle, tendon mass and length). In addition, moment arms of major thoracic limb muscles were measured. Maximum isometric force and power of muscles, the moment of force about a joint, and tendon stress and strain were estimated. Data are compared with those from other cursorial mammals. The thoracic limb of the hare consists predominantly of extrinsic musculature with long parallel fascicles, specialised for generating force over a large range. A large shoulder flexor/elbow extensor muscle mass is present, in particular Triceps brachii. The pennate nature of the long head of this muscle suggests it has an important role in stabilising the elbow joint during stance, whilst moment arm curves suggest that it may also play a role in initiating shoulder flexion. In addition, Supraspinatus and Infraspinatus are capable of generating high forces, potentially to stabilise the shoulder joint during the stance phase of locomotion. Supraspinatus may in addition play an important role in forelimb protraction. The Subscapularis muscle was capable of generating surprisingly high forces, suggesting that the hare must be able to withstand/produce high forces during activities that need medio-lateral stability, such as turning. Distally, tendons were relatively short, showing little potential for elastic energy storage when compared with both their pelvic limb counterparts and their equivalents in the horse thoracic limb. Thus, a 'stiffer' thoracic limb may be beneficial in terms of behaving like a strut, simply supporting and deflecting the body during high-speed running. This more distal/less proximal distribution of limb mass is also likely to be important in retaining the manipulative/adaptive/non-locomotor capabilities of the limb.     

The role of the extrinsic thoracic limb muscles in equine locomotion

Muscles have two major roles in locomotion: to generate force and to absorb/generate power (do work). Economical force generation is achieved by short-fibred pennate muscle while the maximum power output of a muscle is architecture independent. In this study we tested the hypothesis that there is an anatomical and structural separation between the force-generating anti-gravity muscles and the propulsive (limb/trunk moving) muscles of the equine forelimb. Muscle mass and fascicle length measurements were made on the thoracic limb extrinsic muscles of six fresh horse cadavers. Physiological cross-sectional area and maximum isometric force were then estimated. Maximum power was estimated from muscle volume and published contraction velocity data. The majority of extrinsic forelimb muscles were large with long fascicles arranged in parallel to the long axis of the muscle. Muscles arranged in this way are optimised for doing work. The architecture of serratus ventralis thoracis (SVT) was unique. It had short (48 +/- 17 mm) fascicles, arranged at about 45 degrees to the long axis of the muscle, which would suggest a force-generating, anti-gravity role. The muscle belly of SVT was sandwiched between two broad, thick sheets of aponeurosis. Hence, SVT could make a significant contribution to the overall elastic properties of the thoracic limb.     

The role of the extrinsic thoracic limb muscles in equine locomotion

Muscles have two major roles in locomotion: to generate force and to absorb/generate power (do work). Economical force generation is achieved by short-fibred pennate muscle while the maximum power output of a muscle is architecture independent. In this study we tested the hypothesis that there is an anatomical and structural separation between the force-generating anti-gravity muscles and the propulsive (limb/trunk moving) muscles of the equine forelimb. Muscle mass and fascicle length measurements were made on the thoracic limb extrinsic muscles of six fresh horse cadavers. Physiological cross-sectional area and maximum isometric force were then estimated. Maximum power was estimated from muscle volume and published contraction velocity data. The majority of extrinsic forelimb muscles were large with long fascicles arranged in parallel to the long axis of the muscle. Muscles arranged in this way are optimised for doing work. The architecture of serratus ventralis thoracis (SVT) was unique. It had short (48 +/- 17 mm) fascicles, arranged at about 45 degrees to the long axis of the muscle, which would suggest a force-generating, anti-gravity role. The muscle belly of SVT was sandwiched between two broad, thick sheets of aponeurosis. Hence, SVT could make a significant contribution to the overall elastic properties of the thoracic limb.     

Structural and functional anatomy of the neck musculature of the dog (Canis familiaris)

The morphometric properties and the anatomical relationships of the entire musculature of the canine cervical spine are reported herein. These data were obtained from the dissection of cadavers of six dogs. Total muscle length, muscle weight, fascicle length and angles of pennation were recorded for each muscle comprising the canine cervical spine. Based upon these properties, physiological cross-section area (PCSA) and architectural index were estimated. When scaled by whole body mass, the values of each of these parameters were found to be similar between all dogs. Muscles that course from the cranial neck to the shoulder girdle or the rib cage (e.g. brachiocephalicus and rhomboideus capitis) were found to have relatively long fascicles and low PCSA values and thus appear to be designed for rapid excursions. By contrast, muscles that primarily support the neck and shoulder against gravitational forces (e.g. serratus ventralis and trapezius) were found to have relatively high PCSA values and short fascicle lengths, and thus have the capacity to generate large forces. Differences of morphometry as well as nomenclature were found between the canine and human neck musculature. Nevertheless, many similarities exist; in particular, both species have similar muscles adapted to force generation or large excursions. We thus conclude that the canine neck may be used as a modelling tool for biomechanical investigations of the human cervical region as long as the differences listed are borne in mind.     

Behaviour of the human gastrocnemius muscle architecture during submaximal isometric fatigue

The purpose of this study was to examine whether the human gastrocnemius medialis (GM) fascicle length and pennation angle alter during a sustained submaximal isometric plantar flexion. Fourteen male subjects performed maximal voluntary plantar flexions (MVC) on a dynamometer before and after a fatiguing task. This task consisted of a sustained submaximal isometric fatiguing contraction (40% MVC) until failure to hold the defined moment. Ultrasonography was used to visualise the muscle belly of the GM. Leg kinematics were recorded (120 Hz) to calculate the joint moment using inverse dynamics. The exerted moments and the EMG signals from GM and lateralis, soleus and tibialis anterior were measured at 1,080 Hz. The root mean square (RMS) of the EMG signal of the three triceps surae muscles increased significantly (P0.05) between 17% and 28% with fatigue. Further, the fascicle length of the GM significantly decreased from 47.1±8.0 mm at the beginning to 41.8±6.7 mm at the end of fatigue and the pennation angle increased from 23.5±4.1° to 26.3±2.2° (P0.05). The changes in fascicle length and pennation angle of the GM during the contraction can influence the force potential of the muscle due to the force–length relationship and the force transmission to the tendon. This provides evidence on that an additional mechanical mechanism, namely tendon creep, can contribute to the increase in the EMG activity of the GM during submaximal isometric sustained contractions.     

Sonographic studies of human soleus and gastrocnemius muscle architecture: gender variability

The purpose of this study was to establish if there are gender differences in muscle architecture in relaxed human soleus and gastrocnemius muscles of normal, live subjects. Ultrasonography was used to measure fiber bundle length, muscle thickness, and angles of pennation in a total of ten predetermined sites in the medial and lateral heads of gastrocnemius and the anterior and posterior soleus in 19 males and 16 females. Percentage differences between males and females for each parameter were recorded. Gender differences were statistically analyzed using multivariate analysis of variance. In the gastrocnemius and soleus muscles of males and females the differences between the overall fiber bundle length, angle of pennation and muscle thickness were statistically significant (P < 0.05). Overall, females were found to have longer average muscle fiber bundle length and males thicker muscles and larger angles of pennation. The greatest percentage differences of the architectural parameters between males and females were in the posterior soleus: 13% difference in fiber length and 26% difference in angle of pennation in the midline of posterior soleus and 26% difference in muscle thickness of the lateral part of posterior soleus. No correlation was found between leg length and fiber length, angle of pennation or muscle thickness. Fiber length (decreased), angle of pennation (greater) and muscle thickness (greater) of most parts of the gastrocnemius and soleus muscles were significantly different in males and females. Leg length of males and females did not correlate to these architectural parameters. Accepted: 18 February 2000     

Differential adaptations to eccentric versus conventional resistance training in older humans

We hypothesized that training with eccentric contractions only (therefore using higher loads) would yield greater muscle structural and strength gains compared with conventional resistance training. Nine older adults (mean ± s.d. age, 74 ± 3 years) were assigned to a conventional (CONV) resistance training group performing both concentric and eccentric contractions and 10 (age, 67 ± 2 years) to an eccentric-only (ECC) resistance training group. Both groups trained three times per week for 14 weeks at 80% of the five-repetition maximum, specific to each training mode. Maximal knee extensor torque was assessed during isometric, concentric and eccentric contractions across a range of angular velocities (0–3.49 rad s⁻¹). Vastus lateralis muscle architecture (fascicle length, pennation angle and muscle thickness) was assessed in vivo at rest using ultrasonography. Training increased fascicle length in both groups, but the increase was significantly greater in the ECC (20% increase) than the CONV group (8% increase). Conversely, pennation angle significantly increased in the CONV (35% increase) but not in the ECC group (5% increase). Muscle thickness increased to a similar extent in both groups (∼12% increase). In the ECC group, eccentric knee extensor torque increased by 9–17% across velocities, but concentric torque was unchanged. Conversely, in the CONV group, concentric torque increased by 22–37% across velocities, but eccentric torque was unchanged. Instead, isometric torque increased to a similar extent in both groups (∼8% increase). Thus, the two training regimens resulted in differential adaptations in muscle architecture and strength. These results suggest that the stimulus for adding sarcomeres in-series and in-parallel may be different, which implies that different myogenic responses were induced by the two different training methods.     

Inhomogeneous architectural changes of the quadriceps femoris induced by resistance training

Purpose

This study aimed to clarify whether resistance training-induced changes in muscle architecture are homogeneous among the quadriceps and over different regions within each muscle.

Methods

Eleven recreationally active men (27 ± 2 years) completed a 12-week resistance training program for knee extensors. Before and after the intervention, muscle thicknesses, fascicle lengths, and pennation angles of the four muscles (vastus lateralis, vastus medialis, vastus intermedius, rectus femoris) in several regions (2–4 regions per each muscle) were measured using ultrasonography. Anatomical cross-sectional areas (ACSAs) at the same positions as the ultrasound measurements were determined from magnetic resonance images.

Results

Relative increases in the ACSA, muscle thickness, and pennation angle of the rectus femoris were significantly greater than those of the vasti. Relative increases in the ACSAs of the vastus lateralis and rectus femoris were significantly greater in the distal than in the proximal region, and those in the muscle thickness and pennation angle of the vastus intermedius were significantly greater in the medial than in the lateral region. Fascicle lengths did not change in any muscles. The interrelations between muscle thickness and pennation angle remained unchanged after the intervention, with a significant association between the relative changes of the two variables.

Conclusion

The current results indicate that (1) hypertrophy of the quadriceps femoris is associated with a proportional increase in pennation angle but not necessarily in fascicle length, and (2) training-induced changes in muscle size and pennation do not evenly occur among the quadriceps, along or across a muscle.     

Monitoring muscle over three orders of magnitude: Widespread positive allometry among locomotor and body support musculature in the pectoral girdle of varanid lizards (Varanidae)

There is a functional trade-off in the design of skeletal muscle. Muscle strength depends on the number of muscle fibers in parallel, while shortening velocity and operational distance depend on fascicle length, leading to a trade-off between the maximum force a muscle can produce and its ability to change length and contract rapidly. This trade-off becomes even more pronounced as animals increase in size because muscle strength scales with area (length²) while body mass scales with volume (length³). In order to understand this muscle trade-off and how animals deal with the biomechanical consequences of size, we investigated muscle properties in the pectoral girdle of varanid lizards. Varanids are an ideal group to study the scaling of muscle properties because they retain similar body proportions and posture across five orders of magnitude in body mass and are highly active, terrestrially adapted reptiles. We measured muscle mass, physiological cross-sectional area, fascicle length, proximal and distal tendon lengths, and proximal and distal moment arms for 27 pectoral girdle muscles in 13 individuals across 8 species ranging from 64 g to 40 kg. Standard and phylogenetically informed reduced major axis regression was used to investigate how muscle architecture properties scale with body size. Allometric growth was widespread for muscle mass (scaling exponent >1), physiological cross-sectional area (scaling exponent >0.66), but not tendon length (scaling exponent >0.33). Positive allometry for muscle mass was universal among muscles responsible for translating the trunk forward and flexing the elbow, and nearly universal among humeral protractors and wrist flexors. Positive allometry for PCSA was also common among trunk translators and humeral protractors, though less so than muscle mass. Positive scaling for fascicle length was not widespread, but common among humeral protractors. A higher proportion of pectoral girdle muscles scaled with positive allometry than our previous work showed for the pelvic girdle, suggesting that the center of mass may move cranially with body size in varanids, or that the pectoral girdle may assume a more dominant role in locomotion in larger species. Scaling exponents for physiological cross-sectional area among muscles primarily associated with propulsion or with a dual role were generally higher than those associated primarily with support against gravity, suggesting that locomotor demands have at least an equal influence on muscle architecture as body support. Overall, these results suggest that larger varanids compensate for the increased biomechanical demands of locomotion and body support at higher body sizes by developing larger pectoral muscles with higher physiological cross-sectional areas. The isometric scaling rates for fascicle length among locomotion-oriented pectoral girdle muscles suggest that larger varanids may be forced to use shorter stride lengths, but this problem may be circumvented by increases in limb excursion afforded by the sliding coracosternal joint.