Charlotte E. Miller

Ontogenetic scaling of foot musculoskeletal anatomy in elephants

This study quantifies the shape change in elephant manus and pes anatomy with increasing body mass, using computed tomographic scanning. Most manus and pes bones, and manus tendons, maintain their shape, or become more gracile, through ontogeny. Contrary to this, tendons of the pes become significantly more robust, suggesting functional adaptation to increasingly high loads. Ankle tendon cross-sectional area (CSA) scales the highest in the long digital extensor, proportional to body mass1.08±0.21, significantly greater than the highest-scaling wrist tendon (extensor carpi ulnaris, body mass0.69±0.09). These patterns of shape change relate to the marked anatomical differences between the pillar-like manus and tripod-like pes, consistent with differences in fore- and hindlimb locomotor function. The cartilaginous predigits (prepollux and prehallux) of the manus and pes also become relatively more robust through ontogeny, and their pattern of shape change does not resemble that seen in any of the 10 metacarpals and metatarsals. Their CSAs scale above isometry proportional to body mass0.73±0.09 and body mass0.82±0.07 respectively. We infer a supportive function for these structures, preventing collapse of the foot pad during locomotion.

From flat foot to fat foot: structure, ontogeny, function, and evolution of elephant "sixth toes".

Elephants have an extra bone in their feet that can be traced through their fossil record to reveal a transition from flat to raised feet. Several groups of tetrapods have expanded sesamoid (small, tendon-anchoring) bones into digit-like structures (“predigits”), such as pandas’ “thumbs.” Elephants similarly have expanded structures in the fat pads of their fore- and hindfeet, but for three centuries these have been overlooked as mere cartilaginous curiosities. We show that these are indeed massive sesamoids that employ a patchy mode of ossification of a massive cartilaginous precursor and that the predigits act functionally like digits. Further, we reveal clear osteological correlates of predigit joint articulation with the carpals/tarsals that are visible in fossils. Our survey shows that basal proboscideans were relatively “flat-footed” (plantigrade), whereas early elephantiforms evolved the more derived “tip-toed” (subunguligrade) morphology, including the predigits and fat pad, of extant elephants. Thus, elephants co-opted sesamoid bones into a role as false digits and used them for support as they changed their foot posture.

Integration of biomechanical compliance, leverage, and power in elephant limbs

The structure and motion of elephant limbs are unusual compared with those of other animals. Elephants stand and move with straighter limbs (at least when walking), and have limited speed and gait. We devised novel experiments to examine how the limbs of elephants support and propel their mass and to explore the factors that may constrain locomotor performance in these largest of living land animals. We demonstrate that elephant limbs are remarkably compliant even in walking, which maintains low peak forces. Dogma defines elephant limbs as extremely “columnar” for effective weight support, but we demonstrate that limb effective mechanical advantage (EMA) is roughly one-third of that predicted for their size. EMA in elephants is actually smaller than that in horses, which are only one-tenth their mass; it is comparable to human limb values. EMA drops sharply with speed in elephants, as it does in humans. Muscle forces therefore must increase as the limbs become more flexed, and we show how this flexion translates to greater volumes of muscle recruited for locomotion and hence metabolic cost. Surprisingly, elephants use their forelimbs and hindlimbs in similar braking and propulsive roles, not dividing these functions among limbs as was previously assumed or as in other quadrupeds. Thus, their limb function is analogous to four-wheel-drive vehicles. To achieve the observed limb compliance and low peak forces, elephants synchronize their limb dynamics in the vertical direction, but incur considerable mechanical costs from limbs working against each other horizontally.

The Anatomical Foundation for Multidisciplinary Studies of Animal Limb Function: Examples from Dinosaur and Elephant Limb Imaging Studies

What makes so many animals, living and extinct, so popular and distinct is anatomy; it is what leaps out at a viewer first whether they observe a museum’s mounted Tyrannosaurus skeleton or an elephant placidly browsing on the savannah. Anatomy alone can make an animal fascinating — so many animals are so physically unlike human observers, yet what do these anatomical differences mean for the lives of animals?

Pedal Claw Curvature in Birds, Lizards and Mesozoic Dinosaurs – Complicated Categories and Compensating for Mass-Specific and Phylogenetic Control

Pedal claw geometry can be used to predict behaviour in extant tetrapods and has frequently been used as an indicator of lifestyle and ecology in Mesozoic birds and other fossil reptiles, sometimes without acknowledgement of the caveat that data from other aspects of morphology and proportions also need to be considered. Variation in styles of measurement (both inner and outer claw curvature angles) has made it difficult to compare results across studies, as have over-simplified ecological categories. We sought to increase sample size in a new analysis devised to test claw geometry against ecological niche. We found that taxa from different behavioural categories overlapped extensively in claw geometry. Whilst most taxa plotted as predicted, some fossil taxa were recovered in unexpected positions. Inner and outer claw curvatures were statistically correlated, and both correlated with relative claw robusticity (mid-point claw height). We corrected for mass and phylogeny, as both likely influence claw morphology. We conclude that there is no strong mass-specific effect on claw curvature; furthermore, correlations between claw geometry and behaviour are consistent across disparate clades. By using independent contrasts to correct for phylogeny, we found little significant relationship between claw geometry and behaviour. ‘Ground-dweller’ claws are less curved and relatively dorsoventrally deep relative to those of other behavioural categories; beyond this it is difficult to assign an explicit category to a claw based purely on geometry.

An investigation of the dynamic relationship between navicular drop and first metatarsophalangeal joint dorsal excursion

The modern human foot is a complex biomechanical structure that must act both as a shock absorber and as a propulsive strut during the stance phase of gait. Understanding the ways in which foot segments interact can illuminate the mechanics of foot function in healthy and pathological humans. It has been proposed that increased values of medial longitudinal arch deformation can limit metatarsophalangeal joint excursion via tension in the plantar aponeurosis. However, this model has not been tested directly in a dynamic setting. In this study, we tested the hypothesis that during the stance phase, subtalar pronation (stretching of the plantar aponeurosis and subsequent lowering of the medial longitudinal arch) will negatively affect the amount of first metatarsophalangeal joint excursion occurring at push‐off. Vertical descent of the navicular (a proxy for subtalar pronation) and first metatarsophalangeal joint dorsal excursion were measured during steady locomotion over a flat substrate on a novel sample consisting of asymptomatic adult males and females, many of whom are habitually unshod. Least‐squares regression analyses indicated that, contrary to the hypothesis, navicular drop did not explain a significant amount of variation in first metatarsophalangeal joint dorsal excursion. These results suggest that, in an asymptomatic subject, the plantar aponeurosis and the associated foot bones can function effectively within the normal range of subtalar pronation that takes place during walking gait. From a clinical standpoint, this study highlights the need for investigating the in vivo kinematic relationship between subtalar pronation and metatarsophalangeal joint dorsiflexion in symptomatic populations, and also the need to explore other factors that may affect the kinematics of asymptomatic feet.