Metacarpophalangeal joint loads during bonobo locomotion: model predictions versus proxies.

The analysis of internal trabecular and cortical bone has been an informative tool for drawing inferences about behaviour in extant and fossil primate taxa. Within the hand, metacarpal bone architecture has been shown to correlate well with primate locomotion; however, the extent of morphological differences across taxa is unexpectedly small given the variability in hand use. One explanation for this observation is that the activity-related differences in the joint loads acting on the bone are simply smaller than estimated based on commonly used proxies (i.e. external loading and joint posture), which neglect the influence of muscle forces. In this study, experimental data and a musculoskeletal finger model are used to test this hypothesis by comparing differences between climbing and knuckle-walking locomotion of captive bonobos (Pan paniscus) based on (i) joint load magnitude and direction predicted by the models and (ii) proxy estimations. The results showed that the activity-related differences in predicted joint loads are indeed much smaller than the proxies would suggest, with joint load magnitudes being almost identical between the two locomotor modes. Differences in joint load directions were smaller but still evident, indicating that joint load directions might be a more robust indicator of variation in hand use than joint load magnitudes. Overall, this study emphasizes the importance of including muscular forces in the interpretation of skeletal remains and promotes the use of musculoskeletal models for correct functional interpretations.

Musculoskeletal models of a human and bonobo finger: parameter identification and comparison to in vitro experiments.

INTRODUCTION: Knowledge of internal finger loading during human and non-human primate activities such as tool use or knuckle-walking has become increasingly important to reconstruct the behaviour of fossil hominins based on bone morphology. Musculoskeletal models have proven useful for predicting these internal loads during human activities, but load predictions for non-human primate activities are missing due to a lack of suitable finger models. The main goal of this study was to implement both a human and a representative non-human primate finger model to facilitate comparative studies on metacarpal bone loading. To ensure that the model predictions are sufficiently accurate, the specific goals were: (1) to identify species-specific model parameters based on in vitro measured fingertip forces resulting from single tendon loading and (2) to evaluate the model accuracy of predicted fingertip forces and net metacarpal bone loading in a different loading scenario. MATERIALS & METHODS: Three human and one bonobo (Pan paniscus) fingers were tested in vitro using a previously developed experimental setup. The cadaveric fingers were positioned in four static postures and load was applied by attaching weights to the tendons of the finger muscles. For parameter identification, fingertip forces were measured by loading each tendon individually in each posture. For the evaluation of model accuracy, the extrinsic flexor muscles were loaded simultaneously and both the fingertip force and net metacarpal bone force were measured. The finger models were implemented using custom Python scripts. Initial parameters were taken from literature for the human model and own dissection data for the bonobo model. Optimized model parameters were identified by minimizing the error between predicted and experimentally measured fingertip forces. Fingertip forces and net metacarpal bone loading in the combined loading scenario were predicted using the optimized models and the remaining error with respect to the experimental data was evaluated. RESULTS: The parameter identification procedure led to minor model adjustments but considerably reduced the error in the predicted fingertip forces (root mean square error reduced from 0.53/0.69 N to 0.11/0.20 N for the human/bonobo model). Both models remained physiologically plausible after the parameter identification. In the combined loading scenario, fingertip and net metacarpal forces were predicted with average directional errors below 6° and magnitude errors below 12%. CONCLUSIONS: This study presents the first attempt to implement both a human and non-human primate finger model for comparative palaeoanthropological studies. The good agreement between predicted and experimental forces involving the action of extrinsic flexors-which are most relevant for forceful grasping-shows that the models are likely sufficiently accurate for comparisons of internal loads occurring during human and non-human primate manual activities.

Orientation and location of the finite helical axis of the equine forelimb joints.

To reduce anatomically unrealistic limb postures in a virtual musculoskeletal model of a horse's forelimb, accurate knowledge on forelimb joint constraints is essential. The aim of this cadaver study is to report all orientation and position changes of the finite helical axes (FHA) as a function of joint angle for different equine forelimb joints. Five horse cadaver forelimbs with standardized cuts at the midlevel of each segment were used. Bone pins with reflective marker triads were drilled into the forelimb bones. Unless joint angles were anatomically coupled, each joint was manually moved independently in all three rotational degrees of freedom (flexion-extension, abduction-adduction, internal-external rotation). The 3D coordinates of the marker triads were recorded using a six infra-red camera system. The FHA and its orientational and positional properties were calculated and expressed against joint angle over the entire range of motion using a finite helical axis method. When coupled, joint angles and FHA were expressed in function of flexion-extension angle. Flexion-extension movement was substantial in all forelimb joints, the shoulder allowed additional considerable motion in all three rotational degrees of freedoms. The position of the FHA was constant in the fetlock and elbow and a constant orientation of the FHA was found in the shoulder. Orientation and position changes of the FHA over the entire range of motion were observed in the carpus and the interphalangeal joints. We report FHA position and orientation changes as a function of flexion-extension angle to allow for inclusion in a musculoskeletal model of a horse to minimize calculation errors caused by incorrect location of the FHA.

Segmental morphometrics of bonobos (Pan paniscus): are they really different from chimpanzees (Pan troglodytes)?

The inertial properties of body segments reflect performance and locomotor habits in primates. While Pan paniscus is generally described as more gracile, lighter in body mass, and as having relatively longer and heavier hindlimbs than Pan troglodytes, both species exhibit very similar patterns of (quadrupedal and bipedal) kinematics, but show slightly different locomotor repertoires. We used a geometric model to estimate the inertial properties for all body segments (i.e. head, trunk, upper and lower arms, hand, thigh, shank and foot) using external length and diameter measurements of 12 anaesthetized bonobos (eight adults and four immatures). We also calculated whole limb inertial properties. When we compared absolute and relative segment morphometric and inertial variables between bonobos and chimpanzees, we found that adult bonobos are significantly lighter than adult chimpanzees. The bonobo is also shorter in head length, upper and lower arm lengths, and foot length, and is generally lighter in most absolute segment mass values (except head and hand). In contrast, the bonobo has a longer trunk. When scaled relative to body mass, most differences disappear between the two species. Only the longer trunk and the shorter head of the bonobo remain apparent, as well as the lighter thigh compared with the chimpanzee. We found similar values of natural pendular periods of the limbs in both species, despite differences in absolute limb lengths, masses, mass centres (for the hindlimb) and moments of inertia. While our data contradict the commonly accepted view that bonobos have relatively longer and heavier hindlimbs than chimpanzees, they are consistent with the observed similarities in the quadrupedal and bipedal kinematics between these species. The morphological differences between both species are more subtle than those previously described from postcranial osteological materials.

Sagittal plane fore hoof unevenness is associated with fore and hindlimb asymmetrical force vectors in the sagittal and frontal planes.

Asymmetry in forelimb dorsal hoof wall angles, termed unevenness, is associated with forelimb gait asymmetries, but compensatory mechanisms and out of plane ground reaction forces (GRFs) due to unevenness have yet to be documented. The aim of this study was therefore to investigate the effects of fore hoof unevenness on contralateral fore and hind limb force vectors patterns, in both sagittal and frontal planes. A group of n = 34 riding horses were classified into four groups: hoof angle difference of more than 1.5 degrees (UNEVEN; n = 27), including higher left fore (HIGH-LF; n = 12), higher right fore (HIGH-RF; n = 15), and hoof angle difference of less than 1.5 degrees (EVEN; n = 7). Three dimensional ground reaction forces GRFs were collected during trotting. GRF summary vectors representing the magnitude (VecMag) and angular direction (VecAng) of the entire stance phase in the sagittal and the frontal plane were calculated. The effects of unevenness on GRF production were explored using linear regression, repeated measures ANOVA and statistical parametric mapping (SPM) with significance at (P<0.05). In all uneven groups, increasing unevenness affected sagittal VecAng values in the forelimbs, with more propulsive GRF in the high hoof. In the HIGH-RF group, medial GRFs were also found in the high RF hoof compared to lateral GRFs in the low LF hoof (RF VecAng: 0.97±1.64 (deg); LF VecAng: -0.64±1.19 (deg); P<0.05). In both HIGH groups, compensatory associations to increasing unevenness were only found in the RH, but also a significantly greater lateral VecAng was found in the LH (P<0.05) compared to the RH limb. No significant differences (P>0.05) were found between hindlimb pairs in the EVEN group. Unbalanced sagittal and increased frontal plane GRFs in uneven horses suggest that they have greater locomotory challenges, as the equine musculoskeletal system is not constructed to withstand movement and loading in the frontal plane as effectively as it is in the sagittal plane.

Evaluation of a pictorial method to obtain subject-specific inertial properties in equine limb segments.

Data describing segmental masses and moments of inertia (MOI) of limb segments are required for inverse dynamic calculations. In horses, these values are usually calculated using regression equations that have been developed from a limited number of horses representing a small number of breeds. The objective of the present study was to evaluate the performance of a scaling method and a pictorial method for estimating of the values of segmental masses, lengths, and MOI in the equine limb segments by comparing their output with the standard technique involving direct measurements. Limbs of 30 horses of various breeds and sizes were disarticulated post mortem. Segmental masses, lengths, and MOI were determined using a standard method based on weighing the segments, measuring their length with calipers, and estimating the MOI from the rotational frequency of a trifilar pendulum. The scaling method used a jack-knifing procedure to avoid the need for two data sets. The pictorial method was based on digitization of two orthogonal photographs to determine segmental volumes, which were combined with published values for average segment densities to determine the inertial parameters. Scaling method and pictorial method provided comparable estimation of segmental messes and lengths, but the scaling method performed better in estimating segmental MOI. The scaling method worked well enough in the majority of horses but there were a few horses in which it was less effective. The pictorial method sometimes showed a bias correctable by regression equations but it may not warrant the additional effort unless for specific cases.

Hand pressures during arboreal locomotion in captive bonobos (Pan paniscus).

Evolution of the human hand has undergone a transition from use during locomotion to use primarily for manipulation. Previous comparative morphological and biomechanical studies have focused on potential changes in manipulative abilities during human hand evolution, but few have focused on functional signals for arboreal locomotion. Here, we provide this comparative context though the first analysis of hand loading in captive bonobos during arboreal locomotion. We quantify pressure experienced by the fingers, palm and thumb in bonobos during vertical locomotion, suspension and arboreal knuckle-walking. The results show that pressure experienced by the fingers is significantly higher during knuckle-walking compared with similar pressures experienced by the fingers and palm during suspensory and vertical locomotion. Peak pressure is most often experienced at or around the third digit in all locomotor modes. Pressure quantified for the thumb is either very low or absent, despite the thumb making contact with the substrate during all suspensory and vertical locomotor trials. Unlike chimpanzees, bonobos do not show a rolling pattern of digit contact with the substrate during arboreal knuckle-walking - instead, we found that digits 3 and 4 typically touch down first and digit 5 almost always made contact with the substrate. These results have implications for interpreting extant and fossilized hand morphology; we expect bonobo (and chimpanzee) bony morphology to primarily reflect the biomechanical loading of knuckle-walking, while functional signals for arboreal locomotion in fossil hominins are most likely to appear in the fingers, particularly digit 3, and least likely to appear in the morphology of the thumb.

Interspecific variation in the tetradactyl manus of modern tapirs (Perissodactyla: Tapirus) exposed using geometric morphometrics.

The distal forelimb (autopodium) of quadrupedal mammals is a key morphological unit involved in locomotion, body support, and interaction with the substrate. The manus of the tapir (Perissodactyla: Tapirus) is unique within modern perissodactyls, as it retains the plesiomorphic tetradactyl (four-toed) condition also exhibited by basal equids and rhinoceroses. Tapirs are known to exhibit anatomical mesaxonic symmetry in the manus, although interspecific differences and biomechanical mesaxony have yet to be rigorously tested. Here, we investigate variation in the manus morphology of four modern tapir species (Tapirus indicus, Tapirus bairdii, Tapirus pinchaque, and Tapirus terrestris) using a geometric morphometric approach. Autopodial bones were laser scanned to capture surface shape and morphology was quantified using 3D-landmark analysis. Landmarks were aligned using Generalised Procrustes Analysis, with discriminant function and partial least square analyses performed on aligned coordinate data to identify features that significantly separate tapir species. Overall, our results support the previously held hypothesis that T. indicus is morphologically separate from neotropical tapirs; however, previous conclusions regarding function from morphological differences are shown to require reassessment. We find evidence indicating that T. bairdii exhibits reduced reliance on the lateral fifth digit compared to other tapirs. Morphometric assessment of the metacarpophalangeal joint and the morphology of the distal facets of the lunate lend evidence toward high loading on the lateral digits of both the large T. indicus (large body mass) and the small, long limbed T. pinchaque (ground impact). Our results support other recent studies on T. pinchaque, suggesting subtle but important adaptations to a compliant but inclined habitat. In conclusion, we demonstrate further evidence that the modern tapir forelimb is a variable locomotor unit with a range of interspecific features tailored to habitual and biomechanical needs of each species.

A horse's locomotor signature: COP path determined by the individual limb.

INTRODUCTION: Ground reaction forces in sound horses with asymmetric hooves show systematic differences in the horizontal braking force and relative timing of break-over. The Center Of Pressure (COP) path quantifies the dynamic load distribution under the hoof in a moving horse. The objective was to test whether anatomical asymmetry, quantified by the difference in dorsal wall angle between the left and right forelimbs, correlates with asymmetry in the COP path between these limbs. In addition, repeatability of the COP path was investigated. METHODS: A larger group (n = 31) visually sound horses with various degree of dorsal hoof wall asymmetry trotted three times over a pressure mat. COP path was determined in a hoof-bound coordinate system. A relationship between correlations between left and right COP paths and degree of asymmetry was investigated. RESULTS: Using a hoof-bound coordinate system made the COP path highly repeatable and unique for each limb. The craniocaudal patterns are usually highly correlated between left and right, but the mediolateral patterns are not. Some patterns were found between COP path and dorsal wall angle but asymmetry in dorsal wall angle did not necessarily result in asymmetry in COP path and the same could be stated for symmetry. CONCLUSION: This method is a highly sensitive method to quantify the net result of the interaction between all of the forces and torques that occur in the limb and its inertial properties. We argue that changes in motor control, muscle force, inertial properties, kinematics and kinetics can potentially be picked up at an early stage using this method and could therefore be used as an early detection method for changes in the musculoskeletal apparatus.

A three-dimensional morphometric analysis of upper forelimb morphology in the enigmatic tapir (Perissodactyla: Tapirus) hints at subtle variations in locomotor ecology.

Forelimb morphology is an indicator for terrestrial locomotor ecology. The limb morphology of the enigmatic tapir (Perissodactyla: Tapirus) has often been compared to that of basal perissodactyls, despite the lack of quantitative studies comparing forelimb variation in modern tapirs. Here, we present a quantitative assessment of tapir upper forelimb osteology using three-dimensional geometric morphometrics to test whether the four modern tapir species are monomorphic in their forelimb skeleton. The shape of the upper forelimb bones across four species (T. indicus; T. bairdii; T. terrestris; T. pinchaque) was investigated. Bones were laser scanned to capture surface morphology and 3D landmark analysis was used to quantify shape. Discriminant function analyses were performed to reveal features which could be used for interspecific discrimination. Overall our results show that the appendicular skeleton contains notable interspecific differences. We demonstrate that upper forelimb bones can be used to discriminate between species (>91% accuracy), with the scapula proving the most diagnostic bone (100% accuracy). Features that most successfully discriminate between the four species include the placement of the cranial angle of the scapula, depth of the humeral condyle, and the caudal deflection of the olecranon. Previous studies comparing the limbs of T. indicus and T. terrestris are corroborated by our quantitative findings. Moreover, the mountain tapir T. pinchaque consistently exhibited the greatest divergence in morphology from the other three species. Despite previous studies describing tapirs as functionally mediportal in their locomotor style, we find osteological evidence suggesting a spectrum of locomotor adaptations in the tapirs. We conclude that modern tapir forelimbs are neither monomorphic nor are tapirs as conserved in their locomotor habits as previously described. J. Morphol. 277:1469-1485, 2016. © 2016 Wiley Periodicals, Inc.

Gait characteristics and spatio-temporal variables of climbing in bonobos (Pan paniscus).

Although much is known about the terrestrial locomotion of great apes, their arboreal locomotion has been studied less extensively. This study investigates arboreal locomotion in bonobos (Pan paniscus), focusing on the gait characteristics and spatio-temporal variables associated with locomotion on a pole. These features are compared across different substrate inclinations (0°, 30°, 45°, 60°, and 90°), and horizontal quadrupedal walking is compared between an arboreal and a terrestrial substrate. Our results show greater variation in footfall patterns with increasing incline, resulting in more lateral gait sequences. During climbing on arboreal inclines, smaller steps and strides but higher stride frequencies and duty factors are found compared to horizontal arboreal walking. This may facilitate better balance control and dynamic stability on the arboreal substrate. We found no gradual change in spatio-temporal variables with increasing incline; instead, the results for all inclines were clustered together. Bonobos take larger strides at lower stride frequencies and lower duty factors on a horizontal arboreal substrate than on a flat terrestrial substrate. We suggest that these changes are the result of the better grip of the grasping feet on an arboreal substrate. Speed modulation of the spatio-temporal variables is similar across substrate inclinations and between substrate types, suggesting a comparable underlying motor control. Finally, we contrast these variables of arboreal inclined climbing with those of terrestrial bipedal locomotion, and briefly discuss the results with respect to the origin of habitual bipedalism. Am. J. Primatol. 78:1165-1177, 2016. © 2016 Wiley Periodicals, Inc.

Effects of acceleration on gait measures in three horse gaits.

Animals switch gaits according to locomotor speed. In terrestrial locomotion, gaits have been defined according to footfall patterns or differences in center of mass (COM) motion, which characterizes mechanisms that are more general and more predictive than footfall patterns. This has generated different variables designed primarily to evaluate steady-speed locomotion, which is easier to standardize in laboratory conditions. However, in the ecology of an animal, steady-state conditions are rare and the ability to accelerate, decelerate and turn is essential. Currently, there are no data available that have tested whether COM variables can be used in accelerative or decelerative conditions. This study used a data set of kinematics and kinetics of horses using three gaits (walk, trot, canter) to evaluate the effects of acceleration (both positive and negative) on commonly used gait descriptors. The goal was to identify variables that distinguish between gaits both at steady state and during acceleration/deceleration. These variables will either be unaffected by acceleration or affected by it in a predictable way. Congruity, phase shift and COM velocity angle did not distinguish between gaits when the dataset included trials in unsteady conditions. Work (positive and negative) and energy recovery distinguished between gaits and showed a clear relationship with acceleration. Hodographs are interesting graphical representations to study COM mechanics, but they are descriptive rather than quantitative. Force angle, collision angle and collision fraction showed a U-shaped relationship with acceleration and seem promising tools for future research in unsteady conditions.

Functional locomotor consequences of uneven forefeet for trot symmetry in individual riding horses.

Left-right symmetrical distal limb conformation can be an important prerequisite for a successful performance, and it is often hypothesized that asymmetric or uneven feet are important enhancing factors for the development of lameness. On a population level, it has been demonstrated that uneven footed horses are retiring earlier from elite level competition, but the biomechanical consequences are not yet known. The objectives of this study were to compare the functional locomotor asymmetries of horses with uneven to those with even feet. Hoof kinetics and distal limb kinematics were collected from horses (n = 34) at trot. Dorsal hoof wall angle was used to classify horses as even or uneven (<1.5 and >1.5° difference between forefeet respectively) and individual feet as flat (<50°), medium (between 50° and 55°) or upright (>55°). Functional kinetic parameters were compared between even and uneven forefeet using MANOVA followed by ANOVA. The relative influences of differences in hoof angle between the forefeet and of absolute hoof angle on functional parameters were analysed using multiple regression analysis (P<0.05). In horses with uneven feet, the side with the flatter foot showed a significantly larger maximal horizontal braking and vertical ground reaction force, a larger vertical fetlock displacement and a suppler fetlock spring. The foot with a steeper hoof angle was linearly correlated with an earlier braking-propulsion transition. The conformational differences between both forefeet were more important for loading characteristics than the individual foot conformation of each individual horse. The differences in vertical force and braking force between uneven forefeet could imply either an asymmetrical loading pattern without a pathological component or a subclinical lameness as a result of a pathological development in the steeper foot.

Comparison of the head and neck position of elite dressage horses during top-level competitions in 1992 versus 2008.

Among veterinary surgeons, interest has recently increased in the role of the horse's neck as a causative factor in complex locomotor disturbances. Specifically, controversy surrounds the trend for the head to be carried behind the vertical (BHV) in contravention of Fédération Equestre Internationale (FEI) rules. The aim of this study was to determine whether the head angulation of elite dressage horses has changed over the last 25 years, and whether head angulation correlates with the competition score awarded. Head angle was measured from videos recorded during the Grand Prix test at the 1992 Olympic Games and the 2008 World Cup Final, during collected canter (CC), collected trot (CT), passage (Pa), and piaffe (Pi). Head angulations were BHV in CC and CT in both 1992 and 2008. The likelihood of being BHV during Pa or Pi was significantly greater in 2008 than in 1992 (P <0.05). Higher scores correlated significantly with head positions that were further BHV during Pi in 2008 (P <0.05). Head angulations were orientated BHV in all paces in 2008, whereas in 1992 this was only the case for CT and CC. These findings support the hypothesis that, in recent years, FEI dressage judges have not penalised horses for a head position BHV. The findings also support the need for further studies of the effects of head and neck position on the health of horses.

Effect of blindfolding on centre of pressure variables in healthy horses during quiet standing.

In a standing horse the centre of pressure (COP), measured as the resultant vertical ground reaction force (GRF) of all supporting limbs, is adjusted in response to visual, vestibular and proprioceptive information. Stabilographic analysis measures balance by tracking COP movements in the horizontal plane. Loss of visual input affects stability of balance in people and has clinical implications in that instability inherent in some neurological diseases increases with the eyes closed. The objectives of this study were to evaluate the visual contribution to postural stability in horses. The hypothesis was that the magnitude and variability of postural sway variables increases when visual input is removed. Vertical GRFs were measured using two synchronized force plates and COP movements were tracked in 20 horses as they stood without visible movements of the hooves, head or neck. Three trials of 60 s duration were recorded under sighted and blindfolded conditions. Stabilographic variables (craniocaudal and mediolateral COP amplitudes, velocities and mean power frequencies and their within-trial variabilities) were calculated and compared using univariate analysis of variance. Compared with the sighted condition, blindfolding increased the magnitude and the within-trial variability of craniocaudal and mediolateral COP amplitudes and mediolateral COP velocity. The findings indicated that loss of visual input had more effect on the measured COP variables in the time domain (amplitudes, velocities) than in the frequency domain (mean power frequency). The effects of blindfolding on postural stability should be further investigated as part of a diagnostic approach to the evaluation of balance in horses with neurological impairment.

Stride to stride variability in joint angle profiles during transitions from trot to canter in horses.

Spontaneous transitions from anti-phase to in-phase manual coordination are explained in the Haken model that describes the two preferred states as stable regions that work as attractors in a stability landscape. Switching between states coincides with a temporary loss of stability. Coordination variability is believed to be indicative of such a loss of stability. In this study, the hypothesis was tested that an increase in variability in the angle profiles of the joints responsible for the transition will precede the transition. A full gait analysis of four miniature horses transitioning from trot to canter was performed. Joint angle profiles were determined for the joints of all four limbs and were time-normalised to stride duration. Per horse and per stride, the coefficient of variance was calculated as the mean standard deviation of the joint profile over all trials divided by the mean joint angle × 100. As hypothesised, the most proximal limb joints (hip, scapulothoracic, shoulder) followed the predictions to a large extent. The variability of the hip joint angle of the trailing hind limb showed a peak of variability at stride 0; this was quickly reduced after the transition was completed. The detection of this brief perturbation in the hip joint indicates the importance of this joint in the transition process. The hip joint is related to the movements of the limb, pelvis and back, which is one of the main differences between symmetrical and asymmetrical gaits.

Development of postural balance in foals.

This study used stabilographic analysis to measure and describe changes in stability during standing in foals from birth to 5 months of age. Stabilographic analysis was performed on newborn foals immediately after first suckling then daily until 1 week of age, weekly until 1 month of age and monthly until 5 months of age. Ground reaction force (GRF) data were collected for periods of 8s with the foal standing on one or two force plates recording at 1000 Hz. Stabilographic variables describing the amplitude, velocity and frequency of center of pressure (COP) movements were derived from the GRF data. Amplitudes, which were initially larger in the craniocaudal direction, decreased over time in both directions, with craniocaudal amplitude becoming smaller than mediolateral amplitude by 1-2 months of age. At birth, COP velocity was larger in the craniocaudal direction, but decreased rapidly to become smaller than mediolateral velocity by 3 months of age. Mean frequency at birth was higher craniocaudally, but became similar in both directions at 2 months of age. The rapid reductions in craniocaudal amplitude and velocity were thought to reflect improvements in strength and coordination of the flexor/extensor musculature. Newborn foals splay their limbs to compensate for poor control of the abductor/adductor musculature and, after the limbs assumed a vertical posture, mediolateral sway velocity increased.

Relationship between morphological and stabilographic variables in standing horses.

A stabilogram plots movements of the centre of pressure (COP) in the horizontal plane. Derived stabilographic variables quantify postural balance, but it is not known if these variables are size dependent. The aims of this study were to determine which morphological variable was most representative of size, which stabilographic variables were most representative of balance and whether size normalisation improved estimates of postural performance. Croup height (0.93-1.77 m), mass (117-666 kg), base of support (BOS) length (0.74-1.18 m) and BOS width (0.22-0.45 m) were measured in 24 horses. Stabilographic variables describing craniocaudal (CC), mediolateral (ML) and resultant amplitudes, velocities and frequencies of COP motion were measured as the horses stood stationary for 15s with fore and hind hooves on separate force plates (960 Hz). Principal component analysis identified morphological and stabilographic components. Morphological variables were consolidated into a single size component that was represented by body mass. Five stabilographic components explained 91% of the variation in sway patterns and five representative stabilographic variables were identified: CC amplitude, CC velocity, CC frequency, ML amplitude and ML frequency. Mass was correlated with CC velocity and ML frequency, with larger horses having smaller CC velocities and slower ML sway frequencies. When horses were grouped by mass (small horses <400 kg; large horses ≥ 400 kg), the within-group values for CC velocity and ML frequency were no longer correlated with mass.

Spatio-temporal gait characteristics during transitions from trot to canter in horses.

Gaits can be defined based upon specific interlimb coordination patterns characteristic of a limited range of speeds, with one or more defining variables changing discontinuously at a transition. With changing speed, horses perform a repertoire of gaits (walk, trot, canter and gallop), with transitions between them. Knowledge of the series of kinematic events necessary to realize a gait is essential for understanding the proximate mechanisms as well as the control underlying gait transitions. We studied the kinematics of the actual transition from trot to canter in miniature horses. The kinematics were characterized at three different levels: the whole-body level, the spatio-temporal level of the foot falls and the level of basic limb kinematics. This concept represents a hierarchy: the horse's center of mass (COM) moves forward by means of the coordinated action of the limbs and changes in the latter are the result of alterations in the basic limb kinematics. Early and short placement of the fore limb was observed before the dissociation of the footfalls of one of the diagonal limb pairs when entering the canter. Dissociation coincided with increased amplitude and wavelength of the oscillations of the trunk in the sagittal plane. The increased amplitude cannot be explained solely by the passive effects of acceleration or by neck and head movements which are inconsistent with the timing of the transition. We propose that the transition is initiated by the fore limb followed by subsequent changes in the hind limbs in a series of kinematic events that take about 2.5 strides to complete.

Prey handling using whole-body fluid dynamics in batoids.

Fluid flow generated by body movements is a foraging tactic that has been exploited by many benthic species. In this study, the kinematics and hydrodynamics of prey handling behavior in little skates, Leucoraja erinacea, and round stingrays, Urobatis halleri, are compared using kinematics and particle image velocimetry. Both species use the body to form a tent to constrain the prey with the pectoral fin edges pressed against the substrate. Stingrays then elevate the head, which increases the volume between the body and the substrate to generate suction, while maintaining pectoral fin contact with the substrate. Meanwhile, the tip of the rostrum is curled upwards to create an opening where fluid is drawn under the body, functionally analogous to suction-feeding fishes. Skates also rotate the rostrum upwards although with the open rostral sides and the smaller fin area weaker fluid flow is generated. However, skates also use a rostral strike behavior in which the rostrum is rapidly rotated downwards pushing fluid towards the substrate to potentially stun or uncover prey. Thus, both species use the anterior portion of the body to direct fluid flow to handle prey albeit in different ways, which may be explained by differences in morphology. Rostral stiffness and pectoral fin insertion onto the rostrum differ between skates and rays and this corresponds to behavioral differences in prey handling resulting in distinct fluid flow patterns. The flexible muscular rostrum and greater fin area of stingrays allow more extensive use of suction to handle prey while the stiff cartilaginous rostrum of skates lacking extensive fin insertion is used as a paddle to strike prey as well as to clear away sand cover.