Background
Previous analyses of factors influencing footfall timings and gait selection in quadru... more Background Previous analyses of factors influencing footfall timings and gait selection in quadrupeds have focused on the implications for energetic cost or gait mechanics separately. Here we present a model for symmetrical walking gaits in quadrupedal mammals that combines both factors, and aims to predict the substrate contexts in which animals will select certain ranges of footfall timings that (1) minimize energetic cost, (2) minimize rolling and pitching moments, or (3) balance the two. We hypothesize that energy recovery will be a priority on all surfaces, and will be the dominant factor determining footfall timings on flat, ground-like surfaces. The ability to resist pitch and roll, however, will play a larger role in determining footfall choice on narrower and more complex branch-like substrates. As a preliminary test of the expectations of the model, we collected sample data on footfall timings in a primate with relatively high flexibility in footfall timings – the squirrel monkey (Saimiri sciureus) – walking on a flat surface, straight pole, and a pole with laterally-projecting branches to simulate simplified ground and branch substrates. We compare limb phase values on these supports to the expectations of the model.
Results As predicted, walking steps on the flat surface tended towards limb phase values that promote energy exchange. Both pole substrates induced limb phase values predicted to favor reduced pitching and rolling moments.
Conclusions These data provide novel insight into the ways in which animals may choose to adjust their behavior in response to movement on flat versus complex substrates and the competing selective factors that influence footfall timing in mammals. These data further suggest a pathway for future investigations using this perspective.
Objective Pathological gaits have been shown to limit transfer between potential (PE) and kinetic... more Objective Pathological gaits have been shown to limit transfer between potential (PE) and kinetic (KE) energy during walking, which can increase locomotor costs. The purpose of this study was to examine whether energy exchange would be limited in people with knee osteoarthritis (OA). Methods Ground reaction forces during walking were collected from 93 subjects with symptomatic knee OA (self-selected and fast speeds) and 13 healthy controls (self-selected speed) and used to calculate their center of mass (COM) movements, PE and KE relationships, and energy recovery during a stride. Correlations and linear regressions examined the impact of energy fluctuation phase and amplitude, walking velocity, body mass, self-reported pain, and radiographic severity on recovery. Paired t-tests were run to compare energy recovery between cohorts. Results Symptomatic knee OA subjects displayed lower energetic recovery during self-selected walking speeds than healthy controls (p=0.0018). PE and KE ph...
Lumbar vertebral morphology has been used as an indicator of locomotor behavior in living and fos... more Lumbar vertebral morphology has been used as an indicator of locomotor behavior in living and fossil mammals. Rigidity within the lumbar region is thought to be important for increasing overall axial rigidity during various forms of locomotion, including bridging between supports, inverted quadrupedalism, gliding, and flying. However, distinguishing between those behaviors using bony features has been challenging. This study used osteological characters of the lumbar vertebrae to attempt to develop fine-grade functional distinctions among different mammalian species in order to make more complete inferences about how the axial skeleton affects locomotor behavior in extant mammals. These same lumbar characters were measured in two extinct species for which locomotor behaviors are well known, the sloth lemurs (Palaeopropithecus and Babakotia radofilai), in order to further evaluate their locomotor behaviors. Results from a principal components analysis of seven measurements, determine...
Humans stand alone from other primates in that we propel our bodies forward on a relatively stiff... more Humans stand alone from other primates in that we propel our bodies forward on a relatively stiff and arched foot and do so by employing an anatomical arrangement of bones and ligaments in the foot that can operate like a "windlass." This is a significant evolutionary innovation, but it is currently unknown when during hominin evolution this mechanism developed and within what genera or species it originated. The presence of recently discovered fossils along with novel research in the past two decades have improved our understanding of foot mechanics in humans and other apes, making it possible to consider this question more fully. Here we review the main elements thought to be involved in the production of an effective, modern human-like windlass mechanism. These elements are the triceps surae, plantar aponeurosis, medial longitudinal arch, and metatarsophalangeal joints. We discuss what is presently known about the evolution of these features and the challenges associate...
Page 66. 3 The Anatomical Foundation for Multidisciplinary Studies of Animal Limb Function: Examp... more Page 66. 3 The Anatomical Foundation for Multidisciplinary Studies of Animal Limb Function: Examples from Dinosaur and Elephant Limb Imaging Studies John R. Hutchinson1, Charlotte Miller1, Guido Fritsch2, and Thomas ...
Journal of the Royal Society, Interface / the Royal Society, 2008
This study quantifies the shape change in elephant manus and pes anatomy with increasing body mas... more 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 mass(1.08+/-0.21), significantly greater than the highest-scaling wrist tendon (extensor carpi ulnaris, body mass(0.69+/-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...
As the largest extant terrestrial animals, elephants do not trot or gallop but can move smoothly ... more As the largest extant terrestrial animals, elephants do not trot or gallop but can move smoothly to faster speeds without markedly changing their kinematics, yet with a shift from vaulting to bouncing kinetics. To understand this unusual mechanism, we quantified the forelimb and hindlimb motions of eight Asian elephants (Elephas maximus) and seven African elephants (Loxodonta africana). We used 240 Hz motion analysis (tracking 10 joint markers) to measure the flexion/extension angles and angular velocities of the limb segments and joints for 288 strides across an eightfold range of speeds (0.6-4.9 m s(-1)) and a sevenfold range of body mass (521-3684 kg). We show that the columnar limb orientation that elephants supposedly exemplify is an oversimplification--few segments or joints are extremely vertical during weight support (especially at faster speeds), and joint flexion during the swing phase is considerable. The 'inflexible' ankle is shown to have potentially spring-li...
Proceedings of the National Academy of Sciences of the United States of America, Jan 13, 2010
The structure and motion of elephant limbs are unusual compared with those of other animals. Elep... more 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,...
Several groups of tetrapods have expanded sesamoid (small, tendon-anchoring) bones into digit-lik... more 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 ...
Background
Previous analyses of factors influencing footfall timings and gait selection in quadru... more Background Previous analyses of factors influencing footfall timings and gait selection in quadrupeds have focused on the implications for energetic cost or gait mechanics separately. Here we present a model for symmetrical walking gaits in quadrupedal mammals that combines both factors, and aims to predict the substrate contexts in which animals will select certain ranges of footfall timings that (1) minimize energetic cost, (2) minimize rolling and pitching moments, or (3) balance the two. We hypothesize that energy recovery will be a priority on all surfaces, and will be the dominant factor determining footfall timings on flat, ground-like surfaces. The ability to resist pitch and roll, however, will play a larger role in determining footfall choice on narrower and more complex branch-like substrates. As a preliminary test of the expectations of the model, we collected sample data on footfall timings in a primate with relatively high flexibility in footfall timings – the squirrel monkey (Saimiri sciureus) – walking on a flat surface, straight pole, and a pole with laterally-projecting branches to simulate simplified ground and branch substrates. We compare limb phase values on these supports to the expectations of the model.
Results As predicted, walking steps on the flat surface tended towards limb phase values that promote energy exchange. Both pole substrates induced limb phase values predicted to favor reduced pitching and rolling moments.
Conclusions These data provide novel insight into the ways in which animals may choose to adjust their behavior in response to movement on flat versus complex substrates and the competing selective factors that influence footfall timing in mammals. These data further suggest a pathway for future investigations using this perspective.
Objective Pathological gaits have been shown to limit transfer between potential (PE) and kinetic... more Objective Pathological gaits have been shown to limit transfer between potential (PE) and kinetic (KE) energy during walking, which can increase locomotor costs. The purpose of this study was to examine whether energy exchange would be limited in people with knee osteoarthritis (OA). Methods Ground reaction forces during walking were collected from 93 subjects with symptomatic knee OA (self-selected and fast speeds) and 13 healthy controls (self-selected speed) and used to calculate their center of mass (COM) movements, PE and KE relationships, and energy recovery during a stride. Correlations and linear regressions examined the impact of energy fluctuation phase and amplitude, walking velocity, body mass, self-reported pain, and radiographic severity on recovery. Paired t-tests were run to compare energy recovery between cohorts. Results Symptomatic knee OA subjects displayed lower energetic recovery during self-selected walking speeds than healthy controls (p=0.0018). PE and KE ph...
Lumbar vertebral morphology has been used as an indicator of locomotor behavior in living and fos... more Lumbar vertebral morphology has been used as an indicator of locomotor behavior in living and fossil mammals. Rigidity within the lumbar region is thought to be important for increasing overall axial rigidity during various forms of locomotion, including bridging between supports, inverted quadrupedalism, gliding, and flying. However, distinguishing between those behaviors using bony features has been challenging. This study used osteological characters of the lumbar vertebrae to attempt to develop fine-grade functional distinctions among different mammalian species in order to make more complete inferences about how the axial skeleton affects locomotor behavior in extant mammals. These same lumbar characters were measured in two extinct species for which locomotor behaviors are well known, the sloth lemurs (Palaeopropithecus and Babakotia radofilai), in order to further evaluate their locomotor behaviors. Results from a principal components analysis of seven measurements, determine...
Humans stand alone from other primates in that we propel our bodies forward on a relatively stiff... more Humans stand alone from other primates in that we propel our bodies forward on a relatively stiff and arched foot and do so by employing an anatomical arrangement of bones and ligaments in the foot that can operate like a "windlass." This is a significant evolutionary innovation, but it is currently unknown when during hominin evolution this mechanism developed and within what genera or species it originated. The presence of recently discovered fossils along with novel research in the past two decades have improved our understanding of foot mechanics in humans and other apes, making it possible to consider this question more fully. Here we review the main elements thought to be involved in the production of an effective, modern human-like windlass mechanism. These elements are the triceps surae, plantar aponeurosis, medial longitudinal arch, and metatarsophalangeal joints. We discuss what is presently known about the evolution of these features and the challenges associate...
Page 66. 3 The Anatomical Foundation for Multidisciplinary Studies of Animal Limb Function: Examp... more Page 66. 3 The Anatomical Foundation for Multidisciplinary Studies of Animal Limb Function: Examples from Dinosaur and Elephant Limb Imaging Studies John R. Hutchinson1, Charlotte Miller1, Guido Fritsch2, and Thomas ...
Journal of the Royal Society, Interface / the Royal Society, 2008
This study quantifies the shape change in elephant manus and pes anatomy with increasing body mas... more 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 mass(1.08+/-0.21), significantly greater than the highest-scaling wrist tendon (extensor carpi ulnaris, body mass(0.69+/-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...
As the largest extant terrestrial animals, elephants do not trot or gallop but can move smoothly ... more As the largest extant terrestrial animals, elephants do not trot or gallop but can move smoothly to faster speeds without markedly changing their kinematics, yet with a shift from vaulting to bouncing kinetics. To understand this unusual mechanism, we quantified the forelimb and hindlimb motions of eight Asian elephants (Elephas maximus) and seven African elephants (Loxodonta africana). We used 240 Hz motion analysis (tracking 10 joint markers) to measure the flexion/extension angles and angular velocities of the limb segments and joints for 288 strides across an eightfold range of speeds (0.6-4.9 m s(-1)) and a sevenfold range of body mass (521-3684 kg). We show that the columnar limb orientation that elephants supposedly exemplify is an oversimplification--few segments or joints are extremely vertical during weight support (especially at faster speeds), and joint flexion during the swing phase is considerable. The 'inflexible' ankle is shown to have potentially spring-li...
Proceedings of the National Academy of Sciences of the United States of America, Jan 13, 2010
The structure and motion of elephant limbs are unusual compared with those of other animals. Elep... more 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,...
Several groups of tetrapods have expanded sesamoid (small, tendon-anchoring) bones into digit-lik... more 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 ...
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Previous analyses of factors influencing footfall timings and gait selection in quadrupeds have focused on the implications for energetic cost or gait mechanics separately. Here we present a model for symmetrical walking gaits in quadrupedal mammals that combines both factors, and aims to predict the substrate contexts in which animals will select certain ranges of footfall timings that (1) minimize energetic cost, (2) minimize rolling and pitching moments, or (3) balance the two. We hypothesize that energy recovery will be a priority on all surfaces, and will be the dominant factor determining footfall timings on flat, ground-like surfaces. The ability to resist pitch and roll, however, will play a larger role in determining footfall choice on narrower and more complex branch-like substrates. As a preliminary test of the expectations of the model, we collected sample data on footfall timings in a primate with relatively high flexibility in footfall timings – the squirrel monkey (Saimiri sciureus) – walking on a flat surface, straight pole, and a pole with laterally-projecting branches to simulate simplified ground and branch substrates. We compare limb phase values on these supports to the expectations of the model.
Results
As predicted, walking steps on the flat surface tended towards limb phase values that promote energy exchange. Both pole substrates induced limb phase values predicted to favor reduced pitching and rolling moments.
Conclusions
These data provide novel insight into the ways in which animals may choose to adjust their behavior in response to movement on flat versus complex substrates and the competing selective factors that influence footfall timing in mammals. These data further suggest a pathway for future investigations using this perspective.
Previous analyses of factors influencing footfall timings and gait selection in quadrupeds have focused on the implications for energetic cost or gait mechanics separately. Here we present a model for symmetrical walking gaits in quadrupedal mammals that combines both factors, and aims to predict the substrate contexts in which animals will select certain ranges of footfall timings that (1) minimize energetic cost, (2) minimize rolling and pitching moments, or (3) balance the two. We hypothesize that energy recovery will be a priority on all surfaces, and will be the dominant factor determining footfall timings on flat, ground-like surfaces. The ability to resist pitch and roll, however, will play a larger role in determining footfall choice on narrower and more complex branch-like substrates. As a preliminary test of the expectations of the model, we collected sample data on footfall timings in a primate with relatively high flexibility in footfall timings – the squirrel monkey (Saimiri sciureus) – walking on a flat surface, straight pole, and a pole with laterally-projecting branches to simulate simplified ground and branch substrates. We compare limb phase values on these supports to the expectations of the model.
Results
As predicted, walking steps on the flat surface tended towards limb phase values that promote energy exchange. Both pole substrates induced limb phase values predicted to favor reduced pitching and rolling moments.
Conclusions
These data provide novel insight into the ways in which animals may choose to adjust their behavior in response to movement on flat versus complex substrates and the competing selective factors that influence footfall timing in mammals. These data further suggest a pathway for future investigations using this perspective.