Climbing parrots achieve pitch stability using forces and free moments produced by axial-appendicular couples.

During vertical climbing, the gravitational moment tends to pitch the animal's head away from the climbing surface and this may be countered by (1) applying a correcting torque at a discrete contact point, or (2) applying opposing horizontal forces at separate contact points to produce a free moment. We tested these potential strategies in small parrots with an experimental climbing apparatus imitating the fine branches and vines of their natural habitat. The birds climbed on a vertical ladder with four instrumented rungs that measured three-dimensional force and torque, representing the first measurements of multiple contacts from a climbing bird. The parrots ascend primarily by pulling themselves upward using the beak and feet. They resist the gravitational pitching moment with a free moment produced by horizontal force couples between the beak and feet during the first third of the stride and the tail and feet during the last third of the stride. The reaction torque from individual rungs did not counter, but exacerbated the gravitational pitching moment, which was countered entirely by the free moment. Possible climbing limitations were explored using two different rung radii, each with low and high friction surfaces. Rung torque was limited in the large-radius, low-friction condition; however, rung condition did not significantly influence the free moments produced. These findings have implications for our understanding of avian locomotor modules (i.e. coordinated actions of the head-neck, hindlimbs and tail), the use of force couples in vertical locomotion, and the evolution of associated structures.

Great power comes at a high (locomotor) cost: the role of muscle fascicle length in the power versus economy performance trade-off.

Muscle design constraints preclude simultaneous specialization of the vertebrate locomotor system for explosive and economical force generation. The resulting performance trade-off between power and economy has been attributed primarily to individual differences in muscle fiber type composition. While certainly crucial for performance specialization, fiber type likely interacts with muscle architectural parameters, such as fascicle length, to produce this trade-off. Longer fascicles composed of more serial sarcomeres can achieve faster shortening velocities, allowing for greater power production. Long fascicles likely reduce economy, however, because more energy-consuming contractile units are activated for a given force production. We hypothesized that longer fascicles are associated with both increased power production and locomotor cost. In 11 power-trained and 13 endurance-trained recreational athletes, we measured (1) muscle fascicle length via ultrasound in the gastrocnemius lateralis, gastrocnemius medialis and vastus lateralis, (2) maximal power during cycling and countermovement jumps, and (3) running cost of transport. We estimated muscle fiber type non-invasively based on the pedaling rate at which maximal cycling power occurred. As predicted, longer gastrocnemius muscle fascicles were correlated with greater lower-body power production and cost of transport. Multiple regression analyses revealed that variability in maximal power was explained by fiber type (46% for cycling, 24% for jumping) and average fascicle length (20% for cycling, 13% for jumping), while average fascicle length accounted for 15% of the variation in cost of transport. These results suggest that, at least for certain muscles, fascicle length plays an important role in the power versus economy performance trade-off.

Sexual dimorphism in human arm power and force: implications for sexual selection on fighting ability.

Sexual dimorphism often arises from selection on specific musculoskeletal traits that improve male fighting performance. In humans, one common form of fighting includes using the fists as weapons. Here, we tested the hypothesis that selection on male fighting performance has led to the evolution of sexual dimorphism in the musculoskeletal system that powers striking with a fist. We compared male and female arm cranking power output, using it as a proxy for the power production component of striking with a fist. Using backward arm cranking as an unselected control, our results indicate the presence of pronounced male-biased sexual dimorphism in muscle performance for protracting the arm to propel the fist forward. We also compared overhead pulling force between males and females, to test the alternative hypothesis that sexual dimorphism in the upper body of humans is a result of selection on male overhead throwing ability. We found weaker support for this hypothesis, with less pronounced sexual dimorphism in overhead arm pulling force. The results of this study add to a set of recently identified characters indicating that sexual selection on male aggressive performance has played a role in the evolution of the human musculoskeletal system and the evolution of sexual dimorphism in hominins.

Musculoskeletal mass and shape are correlated with competitive ability in male house mice (Mus musculus).

Intense physical competition between males for mating opportunities is widespread among mammals. In such agonistic encounters, males with combinations of morphological, physiological and behavioral characters that allow them to dominate an opponent have greater fitness. However, the specific physical traits associated with competitive ability are poorly understood. Larger body size is often correlated with fitness in mammals. Interestingly, fitness is maximized at intermediate body masses in male house mice (Mus musculus), a species with a polygynous mating system in which males compete physically for access to reproductive resources. Here, we used competition trials in semi-natural, mixed-sex population enclosures to directly measure competitive ability in male house mice based on control of a preferred nesting site. We tested the hypothesis that the musculoskeletal systems of male mice demonstrating high competitive ability are more specialized for competition by comparing the masses of 10 major muscle groups and eight bones as well as a set of 12 skeletal shape indices associated with anatomical specialization for fighting performance in a set of nine winners and 20 losers. Winning males possessed several traits hypothesized to enhance performance in male-male contests: relatively greater mass in several muscle groups and bones of the forelimb and hindlimb and larger scapular surface area. Unexpectedly, no measurements of the head and neck differed significantly between winners and losers. These results identify musculoskeletal traits associated with competitive ability in male house mice and suggest that our current understanding of mammalian fighting performance is incomplete and more nuanced than previously considered.

An outside-the-box activity to demonstrate how humans and animals turn.

Developing hands-on activities that engage and excite K-12 students is critical for stimulating interest in science-based careers. We created an activity for high school students that required them to integrate biology and physics concepts to experience how humans and animals maneuver through their environments (i.e., turning). Understanding how turning works is important because it accounts for up to 50% of daily walking steps and is needed for survival when animals elude predators and capture prey. For this activity, student groups used 2 × 4 lumber, wood screws, and a power drill to build an apparatus that, when connected to the body, altered rotational inertia (object's resistance to change in angular motion, I = mass × radius2). Students navigated through a slalom course with the apparatus (increased radius and rotational inertia) and without the apparatus (mass-matched control). Times to complete the course were compared between trials to determine the influence of rotational inertia on turning performance. Students compiled their data, graphed their results, and found that increased rotational inertia decreased turning performance. Results were connected to sports, rehabilitation, and dinosaur evolution. This activity was implemented during local, regional, national, and international outreach events and adapted for use in undergraduate courses as well (total impact, 250 students). At the end of the activity, students were able to 1) describe whether their results supported their hypothesis; 2) explain how radius influences rotational inertia and turning performance; and 3) apply results to real-world examples. Students and teachers appreciated this "outside-the-box" activity with an engineering twist and found it entertaining.

A disparity between locomotor economy and territory-holding ability in male house mice.

Both economical locomotion and physical fighting are important performance traits to many species because of their direct influence on components of Darwinian fitness. Locomotion represents a substantial portion of the total daily energy budget of many animals. Fighting performance often determines individual reproductive fitness through the means of resource control, social dominance and access to mates. However, phenotypic traits that improve either locomotor economy or fighting ability may diminish performance in the other. Here, we tested for a predicted disparity between locomotor economy and competitive ability in wild-derived house mice (Mus musculus). We used 8 week social competition trials in semi-natural enclosures to directly measure male competitive ability through territorial control and female occupancy within territories. We also measured oxygen consumption during locomotion for each mouse using running trials in an enclosed treadmill and open-flow respirometry. Our results show that territory-holding males have higher absolute and mass-specific oxygen consumption when running (i.e. reduced locomotor economy) compared with males that do not control territories. This relationship was present both before and after 8 week competition trials in semi-natural enclosures. This disparity between physical competitive ability and economical locomotion may impose viability costs on males in species for which competition over mates is common and may constrain the evolution of behavioral and phenotypic diversity, particularly in natural settings with environmental and resource variability.

Fast running restricts evolutionary change of the vertebral column in mammals.

The mammalian vertebral column is highly variable, reflecting adaptations to a wide range of lifestyles, from burrowing in moles to flying in bats. However, in many taxa, the number of trunk vertebrae is surprisingly constant. We argue that this constancy results from strong selection against initial changes of these numbers in fast running and agile mammals, whereas such selection is weak in slower-running, sturdier mammals. The rationale is that changes of the number of trunk vertebrae require homeotic transformations from trunk into sacral vertebrae, or vice versa, and mutations toward such transformations generally produce transitional lumbosacral vertebrae that are incompletely fused to the sacrum. We hypothesize that such incomplete homeotic transformations impair flexibility of the lumbosacral joint and thereby threaten survival in species that depend on axial mobility for speed and agility. Such transformations will only marginally affect performance in slow, sturdy species, so that sufficient individuals with transitional vertebrae survive to allow eventual evolutionary changes of trunk vertebral numbers. We present data on fast and slow carnivores and artiodactyls and on slow afrotherians and monotremes that strongly support this hypothesis. The conclusion is that the selective constraints on the count of trunk vertebrae stem from a combination of developmental and biomechanical constraints.

In vitro strain in human metacarpal bones during striking: testing the pugilism hypothesis of hominin hand evolution.

The hands of hominins (i.e. bipedal apes) are distinguished by skeletal proportions that are known to enhance manual dexterity but also allow the formation of a clenched fist. Because male-male physical competition is important in the mating systems of most species of great apes, including humans, we tested the hypothesis that a clenched fist protects the metacarpal bones from injury by reducing the level of strain during striking. We used cadaver arms to measure in vitro strain in metacarpals during forward strikes with buttressed and unbuttressed fist postures and during side slaps with an open palm. If the protective buttressing hypothesis is correct, the clenched fist posture should substantially reduce strain in the metacarpal bones during striking and therefore reduce the risk of fracture. Recorded strains were significantly higher in strikes in which the hand was secured in unbuttressed and slapping postures than in the fully buttressed posture. Our results suggest that humans can safely strike with 55% more force with a fully buttressed fist than with an unbuttressed fist and with twofold more force with a buttressed fist than with an open-hand slap. Thus, the evolutionary significance of the proportions of the hominin hand may be that these are the proportions that improved manual dexterity while at the same time making it possible for the hand to be used as a club during fighting.

The musculoskeletal system of humans is not tuned to maximize the economy of locomotion.

Humans are known to have energetically optimal walking and running speeds at which the cost to travel a given distance is minimized. We hypothesized that "optimal" walking and running speeds would also exist at the level of individual locomotor muscles. Additionally, because humans are 60-70% more economical when they walk than when they run, we predicted that the different muscles would exhibit a greater degree of tuning to the energetically optimal speed during walking than during running. To test these hypotheses, we used electromyography to measure the activity of 13 muscles of the back and legs over a range of walking and running speeds in human subjects and calculated the cumulative activity required from each muscle to traverse a kilometer. We found that activity of each of these muscles was minimized at specific walking and running speeds but the different muscles were not tuned to a particular speed in either gait. Although humans are clearly highly specialized for terrestrial locomotion compared with other great apes, the results of this study indicate that our locomotor muscles are not tuned to specific walking or running speeds and, therefore, do not maximize the economy of locomotion. This pattern may have evolved in response to selection to broaden the range of sustainable running speeds, to improve performance in motor behaviors not related to endurance locomotion, or in response to selection for both.

Competitive ability in male house mice (Mus musculus): genetic influences.

Conspecifics of many animal species physically compete to gain reproductive resources and thus fitness. Despite the importance of competitive ability across the animal kingdom, specific traits that influence or underpin competitive ability are poorly characterized. Here, we investigate whether there are genetic influences on competitive ability within male house mice. Additionally, we examined if litter demographics (litter size and litter sex ratio) influence competitive ability. We phenotyped two generations for a male's ability to possess a reproductive resource--a prime nesting site--using semi-natural enclosures with mixed sex groupings. We used the "Animal Model" coupled with an extensive pedigree to estimate several genetic parameters. Competitive ability was found to be highly heritable, but only displayed a moderate genetic correlation to body mass. Interestingly, litter sex ratio had a weak negative influence on competitive ability. Litter size had no significant influence on competitive ability. Our study also highlights how much remains unknown about the proximal causes of competitive ability.

Protective buttressing of the human fist and the evolution of hominin hands.

The derived proportions of the human hand may provide supportive buttressing that protects the hand from injury when striking with a fist. Flexion of digits 2-5 results in buttressing of the pads of the distal phalanges against the central palm and the palmar pads of the proximal phalanges. Additionally, adduction of the thenar eminence to abut the dorsal surface of the distal phalanges of digits 2 and 3 locks these digits into a solid configuration that may allow a transfer of energy through the thenar eminence to the wrist. To test the hypothesis of a performance advantage, we measured: (1) the forces and rate of change of acceleration (jerk) from maximum effort strikes of subjects striking with a fist and an open hand; (2) the static stiffness of the second metacarpo-phalangeal (MCP) joint in buttressed and unbuttressed fist postures; and (3) static force transfer from digits 2 and 3 to digit 1 also in buttressed and unbuttressed fist postures. We found that peak forces, force impulses and peak jerk did not differ between the closed fist and open palm strikes. However, the structure of the human fist provides buttressing that increases the stiffness of the second MCP joint by fourfold and, as a result of force transfer through the thenar eminence, more than doubles the ability of the proximal phalanges to transmit 'punching' force. Thus, the proportions of the human hand provide a performance advantage when striking with a fist. We propose that the derived proportions of hominin hands reflect, in part, sexual selection to improve fighting performance.

Axial muscle activation provides stabilization against perturbations while running.

Incidence of traumatic brain injury is an important hazard in sports and recreation. Unexpected (blind-sided) impacts with other players, obstacles, and the ground can be particularly dangerous. We believe this is partially due to the lack of muscular activation which would have otherwise provided protective bracing. In this study participants were asked to run on the treadmill while undergoing perturbations applied at the waist which pulled participants in the fore-aft and lateral directions. To determine the effect of unexpected impacts, participants were given a directional audio-visual warning 0.5 s prior to the perturbation in half of the trials and were unwarned in the other half of the trials. Perturbations were given during the start of the stance phase and during the start of the flight phase to examine two distinct points within the locomotor cycle. Muscle activity was monitored in axial muscles before, during, and after the perturbations were given. We hypothesized that the presence of a warning would allow for voluntary axial muscle activity prior to and during perturbations that would provide bracing of the body, and decreased displacement and acceleration of the head compared to unwarned perturbations. Our results indicate that when a warning is given prior to perturbation, the body was displaced significantly less, and the linear acceleration of the head was also significantly lessened in response to some perturbations. The perturbations given in this study caused significant increases in axial muscle activity compared to activity present during control running. We found evidence that cervical and abdominal muscles increased activity in response to the warning and that typically the warned trials displayed a lower reflexive muscle activity response. Additionally, we found a stronger effect of the warnings on muscle activity within the perturbations given during flight phase than those given at stance phase. Results from this study support the hypothesis that knowledge regarding an impending perturbation is used by the neuromuscular system to activate relevant core musculature and provide bracing to the athlete.

Activity of extrinsic limb muscles in dogs at walk, trot and gallop.

The extrinsic limb muscles perform locomotor work and must adapt their activity to changes in gait and locomotor speed, which can alter the work performed by, and forces transmitted across, the proximal fulcra of the limbs where these muscles operate. We recorded electromyographic activity of 23 extrinsic forelimb and hindlimb muscles and one trunk muscle in dogs while they walked, trotted and galloped on a level treadmill. Muscle activity indicates that the basic functions of the extrinsic limb muscles - protraction, retraction and trunk support - are conserved among gaits. The forelimb retains its strut-like behavior in all gaits, as indicated by both the relative inactivity of the retractor muscles (e.g. the pectoralis profundus and the latissimus dorsi) during stance and the protractor muscles (e.g. the pectoralis superficialis and the omotransversarius) in the first half of stance. The hindlimb functions as a propulsive lever in all gaits, as revealed by the similar timing of activity of retractors (e.g. the biceps femoris and the gluteus medius) during stance. Excitation increased in many hindlimb muscles in the order walk-trot-gallop, consistent with greater propulsive impulses in faster gaits. Many forelimb muscles, in contrast, showed the greatest excitation at trot, in accord with a shorter limb oscillation period, greater locomotor work performed by the forelimb and presumably greater absorption of collisional energy.

Precocial hindlimbs and altricial forelimbs: partitioning ontogenetic strategies in mallards (Anas platyrhynchos).

Precocial development, in which juveniles are relatively mature at hatching or birth, is more common among vertebrates than altricial development, and is likely to be the basal condition. Altricial development characterizes many birds and mammals and is generally viewed as an alternate strategy, promoting fast growth rates, short developmental periods and relatively poor locomotor performance prior to attaining adult size. Many aquatic birds such as Anseriformes (ducks, geese and swans), Charadriformes (gulls and terns) and Gruiformes (rails) undergo distinctive developmental trajectories, in that hatchlings are able to run and swim the day they hatch, yet they do not begin to fly until fully grown. We hypothesized that there should be tradeoffs in apportioning bone and muscle mass to the hindlimb and forelimb that could account for these patterns in locomotor behavior within the mallard (Anas platyrhynchos). Growth of the musculoskeletal system in the forelimbs and hindlimbs was measured and compared with maximal aquatic and terrestrial sprint speeds and aerial descent rates throughout the 2-month-long mallard ontogenetic period. At 30 days post hatching, when body mass is 50% of adult values, hindlimb muscle mass averages 90% and forelimb muscle mass averages 10% of adult values; similarly, bone growth (length and width) in the hindlimbs and forelimbs averages 90 and 60% of adult values, respectively. The attainment of mallard locomotor performance parallels the morphological maturation of forelimb and hindlimb morphometrics - hindlimb performance initiates just after hatching at a relatively high level (~50% adult values) and gradually improves throughout the first month of development, while forelimb performance is relatively non-existent at hatching (~10% adult values), experiencing delayed and dramatic improvement in function, and maturing at the time of fledging. This divergence in ontogenetic strategy between locomotor modules could allow developing Anseriformes to inhabit aquatic, predator-reduced refuges without relying on flight for juvenile escape. Furthermore, by freeing the forelimbs from locomotor demand early in ontogeny, Anseriformes may bypass the potential canalization (i.e. retention) of juvenile form present within their precocial hindlimbs, to dramatically depart in forelimb form and function in the adult.

The Human Neck is Part of the Musculoskeletal Core: Cervical Muscles Help Stabilize the Pelvis During Running and Jumping.

During locomotion, cervical muscles must be active to stabilize the head as the body accelerates and decelerates. We hypothesized that cervical muscles are also part of the linked chain of axial muscles that provide core stabilization against torques applied to the hip joint by the extrinsic muscles of the legs. To test whether specific cervical muscles play a role in postural stabilization of the head and/or core stabilization of the pelvic girdle, we used surface electromyography to measure changes in muscle activity in response to force manipulations during constant speed running and maximum effort counter-movement jumps. We found that doubling the mass of the head during both running and maximum effort jumping had little or no effect on (1) acceleration of the body and (2) cervical muscle activity. Application of horizontal forward and rearward directed forces at the pelvis during running tripled mean fore and aft accelerations, thereby increasing both the pitching moments on the head and flexion and extension torques applied to the hip. These manipulations primarily resulted in increases in cervical muscle activity that is appropriate for core stabilization of the pelvis. Additionally, when subjects jumped maximally with an applied downward directed force that reduced acceleration and therefore need for cervical muscles to stabilize the head, cervical muscle activity did not decrease. These results suggest that during locomotion, rather than acting to stabilize the head against the effects of inertia, the superficial muscles of the neck monitored in this study help to stabilize the pelvis against torques imposed by the extrinsic muscles of the legs at the hip joint. We suggest that a division of labor may exist between deep cervical muscles that presumably provide postural stabilization of the head versus superficial cervical muscles that provide core stabilization against torques applied to the pelvic and pectoral girdles by the extrinsic appendicular muscles.

Durante la locomoción, los músculos cervicales deben estar activos para estabilizar la cabeza a medida que el cuerpo acelera y desacelera. Presumimos que los músculos cervicales también son parte de la cadena unida de músculos axiales que brindan estabilización central contra las torsiones aplicadas a la articulación de la cadera por los músculos extrínsecos de las piernas. Para evaluar si los músculos cervicales específicos desempeñan un papel en la estabilización postural de la cabeza y/o la estabilización central de la cintura pélvica, utilizamos electromiografía de superficie para medir los cambios en la actividad muscular en respuesta a las manipulaciones de fuerza durante la carrera a velocidad constante y los saltos con contramovimiento de esfuerzo máximo. Descubrimos que duplicar la masa de la cabeza durante la carrera y el salto de máximo esfuerzo tuvo poco o ningún efecto sobre (1) la aceleración del cuerpo y (2) la actividad de los músculos cervicales. La aplicación de fuerzas horizontales dirigidas hacia adelante y hacia atrás en la pelvis durante la carrera triplicó las aceleraciones medias hacia adelante y hacia atrás, aumentando así tanto los momentos de cabeceo en la cabeza como los pares de flexión y extensión aplicados a la cadera. Estas manipulaciones dieron como resultado principalmente aumentos en la actividad de los músculos cervicales que son apropiados para la estabilización central de la pelvis. Además, cuando los sujetos saltaban al máximo aplicando una fuerza dirigida hacia abajo que reducía la aceleración y, por lo tanto, la necesidad de los músculos cervicales para estabilizar la cabeza, la actividad de los músculos cervicales no disminuía. Sugerimos que puede existir una división del trabajo entre los músculos cervicales profundos que presumiblemente brindan estabilización postural de la cabeza versus los músculos cervicales superficiales que brindan estabilización central contra los torques aplicados a las cinturas pélvica y pectoral por los músculos apendiculares extrínsecos.

Pendant la locomotion, les muscles cervicaux doivent être actifs pour stabiliser la tête lorsque le corps accélère et décélère. Nous avons émis l"hypothèse que les muscles cervicaux font également partie de la chaîne liée des muscles axiaux qui assurent la stabilisation du noyau contre les couples appliqués à l"articulation de la hanche par les muscles extrinsèques des jambes. Pour tester si des muscles cervicaux spécifiques jouent un rôle dans la stabilisation posturale de la tête et/ou la stabilisation centrale de la ceinture pelvienne, nous avons utilisé l"électromyographie de surface pour mesurer les changements dans l"activité musculaire en réponse aux manipulations de force pendant la course à vitesse constante et les sauts de contre-mouvement à effort maximal. Nous avons constaté que doubler la masse de la tête pendant la course et le saut à effort maximal avait peu ou pas d"effet sur (1) l"accélération du corps et (2) l"activité des muscles cervicaux. L"application de forces horizontales dirigées vers l"avant et vers l"arrière au niveau du bassin pendant la course a triplé les accélérations moyennes vers l"avant et vers l"arrière, augmentant ainsi à la fois les moments de tangage sur la tête et les couples de flexion et d"extension appliqués à la hanche. Ces manipulations ont principalement entraîné une augmentation de l"activité des muscles cervicaux, ce qui est approprié pour la stabilisation centrale du bassin. De plus, lorsque les sujets sautaient au maximum avec une force dirigée vers le bas qui réduisait l"accélération et donc le besoin de muscles cervicaux pour stabiliser la tête, l"activité des muscles cervicaux ne diminuait pas. Ces résultats suggèrent que lors de la locomotion, plutôt que d"agir pour stabiliser la tête contre les effets de l"inertie, les muscles superficiels du cou suivis dans cette étude aident à stabiliser le bassin contre les couples imposés par les muscles extrinsèques des jambes au niveau de l"articulation de la hanche. Nous suggérons qu"une division du travail peut exister entre les muscles cervicaux profonds qui assurent vraisemblablement la stabilisation posturale de la tête et les muscles cervicaux superficiels qui assurent la stabilisation centrale contre les couples appliqués aux ceintures pelvienne et pectorale par les muscles appendiculaires extrinsèques.

Effects of fore-aft body mass distribution on acceleration in dogs.

The ability of a quadruped to apply propulsive ground reaction forces (GRF) during rapid acceleration may be limited by muscle power, foot traction or the ability to counteract the nose-up pitching moment due to acceleration. Because the biomechanics of acceleration change, both throughout the stride cycle and over subsequent strides as velocity increases, the factors limiting propulsive force production may also change. Depending on which factors are limiting during each step, alterations in fore-aft body mass distribution may either increase or decrease the maximum propulsive GRF produced. We analyzed the effects of experimental alterations in the fore-aft body mass distribution of dogs as they performed rapid accelerations. We measured the changes in trunk kinematics and GRF as dogs accelerated while carrying 10% body mass in saddlebags positioned just in front of the shoulder girdle or directly over the pelvic girdle. We found that dogs applied greater propulsive forces in the initial hindlimb push-off and first step by the lead forelimb in both weighted conditions. During these steps dogs appear to have been limited by foot traction. For the trailing forelimb, propulsive forces and impulses were reduced when dogs wore caudally placed weights and increased when dogs wore cranially placed weights. This is consistent with nose-up pitching or avoidance thereof having limited propulsive force production by the trailing forelimb. By the second stride, the hindlimbs appear to have been limited by muscle power in their ability to apply propulsive force. Adding weights decreased the propulsive force they applied most in the beginning of stance, when limb retractor muscles were active in supporting body weight. These results suggest that all three factors: foot traction, pitching of the body, and muscle power play roles in limiting quadrupedal acceleration. Digging in to the substrate with claws or hooves appears to be necessary for maximizing propulsion in the initial hindlimb push-off and first forelimb step. Shifting the center of mass forward, as occurred with the loss of the large and heavy tail in the evolution of mammals, is likely to increase the contribution of the forelimbs to acceleration. Hindlimb muscle power appears to play a greater role in limiting acceleration than does forelimb muscle power. As such, we might expect animals built for rapid acceleration to have an increased ratio of hindlimb to forelimb muscle mass.

Function of the epaxial muscles in walking, trotting and galloping dogs: implications for the evolution of epaxial muscle function in tetrapods.

The body axis plays a central role in tetrapod locomotion. It contributes to the work of locomotion, provides the foundation for the production of mechanical work by the limbs, is central to the control of body posture, and integrates limb and trunk actions. The epaxial muscles of mammals have been suggested to mobilize and globally stabilize the trunk, but the timing and the degree to which they serve a particular function likely depend on the gait and the vertebral level. To increase our understanding of their function, we recorded the activity of the m. multifidus lumborum and the m. longissimus thoracis et lumborum at three cranio-caudal levels in dogs while they walked, trotted and galloped. The level of muscle recruitment was significantly higher during trotting than during walking, but was similar during trotting and galloping. During walking, epaxial muscle activity is appropriate to produce lateral bending and resist long-axis torsion of the trunk and forces produced by extrinsic limb muscles. During trotting, they also stabilize the trunk in the sagittal plane against the inertia of the center of mass. Muscle recruitment during galloping is consistent with the production of sagittal extension. The sequential activation along the trunk during walking and galloping is in accord with the previously observed traveling waves of lateral and sagittal bending, respectively, while synchronized activity during trotting is consistent with a standing wave of trunk bending. Thus, the cranio-caudal recruitment patterns observed in dogs resemble plesiomorphic motor patterns of tetrapods. In contrast to other tetrapods, mammals display bilateral activity during symmetrical gaits that provides increased sagittal stability and is related to the evolution of a parasagittal limb posture and greater sagittal mobility.

The cost of ventilation in birds measured via unidirectional artificial ventilation.

The highly derived mechanism birds use to ventilate their lungs relies on dorsoventral excursions of their heavily muscled sternum and abdominal viscera. Our expectation of the level of mechanical work involved in this mechanism led us to hypothesize that the metabolic cost of breathing is higher in birds than in other tetrapods. To test this theory, we used unidirectional artificial ventilation (UDV) to stop normal ventilatory movements in guinea fowl (Numida meleagris L.) at rest and during treadmill locomotion at three speeds. Oxygen consumption was measured during normal breathing and UDV, and the difference was used to approximate the cost of ventilation. Contrary to our prediction, metabolism increased when ventilatory movements ceased during UDV at rest. Although we do not understand why this occurred we suspect that UDV induced a homeostatic mechanism to counteract the loss of carbon dioxide. Nevertheless, across all running speeds, metabolism decreased significantly with UDV, indicating a minimum cost of ventilation during running of 1.43+/-0.62% of total running metabolism or 0.48+/-0.21 mL O(2) (L ventilated)(-1). These results suggest that the metabolic cost of ventilation is low in birds and that it is within the range of costs reported previously for other amniotes.

Sexual dimorphism in postcranial skeletal shape suggests male-biased specialization for physical competition in anthropoid primates.

Sexual dimorphism often arises as a response to selection on traits that improve a male's ability to physically compete for access to mates. In primates, sexual dimorphism in body mass and canine size is more common in species with intense male-male competition. However, in addition to these traits, other musculoskeletal adaptations may improve male fighting performance. Postcranial traits that increase strength, agility, and maneuverability may also be under selection. To test the hypothesis that males, as compared to females, are more specialized for physical competition in their postcranial anatomy, we compared sex-specific skeletal shape using a set of functional indices predicted to improve fighting performance. Across species, we found significant sexual dimorphism in a subset of these indices, indicating the presence of skeletal shape sexual dimorphism in our sample of anthropoid primates. Mean skeletal shape sexual dimorphism was positively correlated with sexual dimorphism in body size, an indicator of the intensity of male-male competition, even when controlling for both body mass and phylogenetic relatedness. These results suggest that selection on male fighting ability has played a role in the evolution of postcranial sexual dimorphism in primates.