No abstract

Scratch-digging mammals are commonly described as having large, powerful forelimb muscles for applying high force to excavate earth, yet studies quantifying the architectural properties of the musculature are largely unavailable. To further test hypotheses about traits that represent specializations for scratch-digging, we quantified muscle architectural properties and myosin expression in the forelimb of the groundhog (Marmota monax), a digger that constructs semi-complex burrows. Architectural properties measured were muscle moment arm, muscle mass (MM), belly length (ML), fascicle length (l F ), pennation angle and physiological cross-sectional area (PCSA), and these metrics were used to estimate maximum isometric force, joint torque and power. Myosin heavy chain (MHC) isoform composition was determined in selected forelimb muscles by SDS-PAGE and densitometry analysis. Groundhogs have large limb retractors and elbow extensors that are capable of applying moderately high torque at the shoulder and elbow joints, respectively. Most of these muscles (e.g. latissimus dorsi and pectoralis superficialis) have high l F /ML ratios, indicating substantial shortening ability and moderate power. The unipennate triceps brachii long head has the largest PCSA and is capable of the highest joint torque at both the shoulder and elbow joints. The carpal and digital flexors show greater pennation and shorter fascicle lengths than the limb retractors and elbow extensors, resulting in higher PCSA/MM ratios and force production capacity. Moreover, the digital flexors have the capacity for both appreciable fascicle shortening and force production, indicating high muscle work potential. Overall, the forelimb musculature of the groundhog is capable of relatively low sustained force and power, and these properties are consistent with the findings of a predominant expression of the MHC-2A isoform. Aside from the apparent modifications to the digital flexors, the collective muscle properties observed are consistent with its behavioral classification as a lessspecialized burrower and these may be more representative of traits common to numerous rodents with burrowing habits or mammals with some fossorial ability.

Moles have modified thoracic limbs with hypertrophied pectoral girdle muscles that allow them to apply remarkably high lateral out-forces during the power stroke when burrowing. To further understand the high force capabilities of mole forelimbs, architectural properties of the thoracic limb muscles were quantified in the Eastern mole (Scalopus aquaticus). Architectural properties measured included muscle mass, moment arm, belly length, fascicle length, and pennation angle, and these were used to provide estimates of maximum isometric force, joint torque, and power. Measurements of muscle moment arms and limb lever lengths were additionally used to analyze the out-force contributions of the major pectoral girdle muscles. Most muscles have relatively long fascicles and little-to-no pennation. The humeral abductor/rotators as a functional group are massive and are capable of relatively high force, power, and joint torque. Of this group, the bipennate m. teres major is the most massive and has the capacity to produce the highest force and joint torque to abduct and axially rotate the humerus. In general, the distal limb muscles are relatively small, but have the capacity for high force and mechanical work by fascicle shortening. The muscle architectural properties of the elbow extensors (e.g., m. triceps brachii) and carpal flexors (e.g., m. palmaris longus) are consistent with the function of these muscles to augment lateral out-force application. The humeral abductor/rotators m. latissimus dorsi, m. teres major, m. pectoralis, and m. subscapularis are calculated to contribute 13.9 N to out-force during the power stroke, and this force is applied in a 'frontal' plane causing abduction of the humerus about the sternoclavicular joint. Moles have several specializations of their digging apparatus that greatly enhance the application of out-force, and these morphological features suggest convergence on limb form and burrowing function between New and Old World moles.

Due to small body size, an immature musculoskeletal system, and other growth-related limits on performance, juvenile mammals frequently experience a greater risk of predation than their adult counterparts. As a result, behaviorally precocious juveniles are hypothesized to exhibit musculoskeletal advantages that permit them to accelerate rapidly and evade predation. This hypothesis was tested through detailed quantitative evaluation of muscle growth in wild Eastern cottontail rabbits (Sylvilagus floridanus). Cottontail rabbits experience high rates of mortality during the first year of life, suggesting that selection might act to improve performance in growing juveniles. Therefore, it was predicted that muscle properties associated with force and power capacity should be enhanced in juvenile rabbits to facilitate enhanced locomotor performance. We quantified muscle architecture from 24 paravertebral and hindlimb muscles across ontogeny in a sample of n = 29 rabbits and evaluated the body mass scaling of muscle mass (MM), physiological cross-sectional area (PCSA), isometric force (F max ), and instantaneous power (P inst ), along with several dimensionless architectural indices. In contrast to our hypothesis, MM and PCSA for most muscles change with positive allometry during growth by scaling at M 1:3 b and M 1:1 b , respectively, whereas F max and P inst generally scale indistinguishably from isometry, as do the architectural indices tested. However, scaling patterns indicate that the digital flexors and ankle extensors of juvenile S. floridanus have greater capacities for force and power, respectively, than those in adults, suggesting these muscle properties may be a part of several compensatory features that promote enhanced acceleration performance in young rabbits. Overall, our study implies that body size constraints place larger, more mature rabbits at a disadvantage during acceleration, and that adults must develop hypertrophied muscles in order to maintain mechanical similarity in force and power capacities across development. These findings challenge the accepted understanding that juvenile animals are at a performance detriment relative to adults. Instead, for prey-predator interactions necessitating short intervals of high force and power generation relative to body mass, as demonstrated by rapid acceleration of cottontail rabbits fleeing predators, it may be the adults that struggle to keep pace with juveniles.