Russell Savage

New insights into the plantar pressure correlates of walking speed using pedobarographic statistical parametric mapping (pSPM)

This study investigates the relation between walking speed and the distribution of peak plantar pressure and compares a traditional ten-region subsampling (10RS) technique with a new technique: pedobarographic statistical parametric mapping (pSPM). Adapted from cerebral fMRI methodology, pSPM is a digital image processing technique that registers foot pressure images such that homologous structures optimally overlap, thereby enabling statistical tests to be conducted at the pixel level. Following previous experimental protocols, we collected pedobarographic records from 10 subjects walking at three different speeds: slow, normal, and fast. Walking speed was recorded and correlated with the peak pressures extracted from the 10 regions, and subsequently with the peak pixel data extracted after pSPM preprocessing. Both methods revealed significant positive correlation between peak plantar pressure and walking speed over the rearfoot and distal forefoot after Bonferroni correction for multiple comparisons. The 10RS analysis found positive correlation in the midfoot and medial proximal forefoot, but the pixel data exhibited significant negative correlation throughout these regions (p<5x10(-5)). Comparing the statistical maps from the two approaches shows that subsampling may conflate pressure differences evident in pixel-level data, obscuring or even reversing statistical trends. The negative correlation observed in the midfoot implies reduced longitudinal arch collapse with higher walking speeds. We infer that this results from pre- or early-stance phase muscle activity and speculate that preferred walking speed reflects, in part, a balance between the energy required to tighten the longitudinal arch and the apparent propulsive benefits of the stiffened arch.

Human-like external function of the foot, and fully upright gait, confirmed in the 3.66 million year old Laetoli hominin footprints by topographic statistics, experimental footprint-formation and computer simulation

It is commonly held that the major functional features of the human foot (e.g. a functional longitudinal medial arch, lateral to medial force transfer and hallucal (big-toe) push-off) appear only in the last 2 Myr, but functional interpretations of footbones and footprints of early human ancestors (hominins) prior to 2 million years ago (Mya) remain contradictory. Pixel-wise topographical statistical analysis of Laetoli footprint morphology, compared with results from experimental studies of footprint formation; foot-pressure measurements in bipedalism of humans and non-human great apes; and computer simulation techniques, indicate that most of these functional features were already present, albeit less strongly expressed than in ourselves, in the maker of the Laetoli G-1 footprint trail, 3.66 Mya. This finding provides strong support to those previous studies which have interpreted the G-1 prints as generally modern in aspect.

Morphological analysis of the hindlimb in apes and humans. II. Moment arms

Flexion/extension moment arms were obtained for the major muscles crossing the hip, knee and ankle joints in the orang‐utan, gibbon, gorilla (Eastern and Western lowland) and bonobo. Moment arms varied with joint motion and were generally longer in proximal limb muscles than distal limb muscles. The shape of the moment arm curves (i.e. the plots of moment arm against joint angle) differed in different hindlimb muscles and in the same muscle in different subjects (both in the same and in different ape species). Most moment arms increased with increasing joint flexion, a finding which may be understood in the context of the employment of flexed postures by most non‐human apes (except orang‐utans) during both terrestrial and arboreal locomotion. When compared with humans, non‐human great apes tended to have muscles better designed for moving the joints through large ranges. This was particularly true of the pedal digital flexors in orang‐utans. In gibbons, the only lesser ape studied here, many of the moment arms measured were relatively short compared with those of great apes. This study was performed on a small sample of apes and thus differences noted here warrant further investigation in larger populations.

Dynamics of longitudinal arch support in relation to walking speed: contribution of the plantar aponeurosis

The plantar aponeurosis (PA), in spanning the whole length of the plantar aspect of the foot, is clearly identified as one of the key structures that is likely to affect compliance and stability of the longitudinal arch. A recent study performed in our laboratory showed that tension/elongation in the PA can be predicted from the kinematics of the segments to which the PA is attached. In the present investigation, stereophotogrammetry and inverse kinematics were employed to shed light on the mechanics of the longitudinal arch and its main passive stabilizer, the PA, in relation to walking speed. When compared with a neutral unloaded position, the medial longitudinal arch underwent greater collapse during the weight‐acceptance phase of stance at higher walking speed (0.1°±1.9° in slow walking; 0.9°±2.6° in fast walking; P = 0.0368). During late stance the arch was higher (3.4°±3.1° in slow walking; 2.8°±2.7° in fast walking; P = 0.0227) and the metatarsophalangeal joints more dorsiflexed (e.g. at the first metatarsophalangeal joint, 52°±5° in slow walking; 64°±4° in fast walking; P < 0.001) during fast walking. Early‐stance tension in the PA increased with speed, whereas maximum tension during late stance did not seem to be significantly affected by walking speed. Although, on the one hand, these results give evidence for the existence of a pre‐heel‐strike, speed‐dependent, arch‐stiffening mechanism, on the other hand they suggest that augmentation of arch height in late stance is enhanced by higher forces exerted by the intrinsic muscles on the plantar aspect of the foot when walking at faster speeds.

The evolution of compliance in the human lateral mid-foot

Fossil evidence for longitudinal arches in the foot is frequently used to constrain the origins of terrestrial bipedality in human ancestors. This approach rests on the prevailing concept that human feet are unique in functioning with a relatively stiff lateral mid-foot, lacking the significant flexion and high plantar pressures present in non-human apes. This paradigm has stood for more than 70 years but has yet to be tested objectively with quantitative data. Herein, we show that plantar pressure records with elevated lateral mid-foot pressures occur frequently in healthy, habitually shod humans, with magnitudes in some individuals approaching absolute maxima across the foot. Furthermore, the same astonishing pressure range is present in bonobos and the orangutan (the most arboreal great ape), yielding overlap with human pressures. Thus, while the mean tendency of habitual mechanics of the mid-foot in healthy humans is indeed consistent with the traditional concept of the lateral mid-foot as a relatively rigid or stabilized structure, it is clear that lateral arch stabilization in humans is not obligate and is often transient. These findings suggest a level of detachment between foot stiffness during gait and osteological structure, hence fossilized bone morphology by itself may only provide a crude indication of mid-foot function in extinct hominins. Evidence for thick plantar tissues in Ardipithecus ramidus suggests that a human-like combination of active and passive modulation of foot compliance by soft tissues extends back into an arboreal context, supporting an arboreal origin of hominin bipedalism in compressive orthogrady. We propose that the musculoskeletal conformation of the modern human mid-foot evolved under selection for a functionally tuneable, rather than obligatory stiff structure.

Inertial properties of hominoid limb segments

Quantitative, accurate data regarding the inertial properties of body segments are of paramount importance when developing musculo‐skeletal locomotor models of living animals and, by inference, their ancestors. The limited number of available primate cadavers, and the destructive nature of the post‐mortem, result in such data being very rare for primates. This study builds on the work of Crompton et al. (Am. J. Phys. Anthropol. 1996, 99, 547–570) and reports inertial properties of the body segments of gorillas, chimpanzees, orang‐utans and gibbons. Segment mass, centre of mass and the radius of gyration of five ape cadavers were measured using a complex‐pendulum technique and compared with the results derived from external measurements of segment lengths and diameters on the same animals. With additional data from external measurements of eight more hominoid cadavers, and published data, intergeneric differences between the inertial properties and the distribution of mass between limb segments are analysed and related to the locomotor habits of the species. We found that segment inertial properties show extensive overlap between ape genera as a result of large interindividual variation. Segment mass distribution also overlaps between apes and humans, with the exception of the shank segment. However, owing to a different distribution of mass between the limb segments, the centre of mass of both the arms and the legs is located more distally in apes than in humans, and the natural pendular period of ape forelimbs is larger than that of the hindlimbs. This suggests that, in contrast to the limbs of cursorial mammals and cercopithecoid primates, hominoid limbs are not optimized for efficiency in quadrupedal walking, but rather reflect a compromise between various locomotor modes. Common chimpanzees may have secondarily evolved a more efficient quadrupedal gait.

Functional adaptations in the forelimb muscles of non-human great apes

The maximum capability of a muscle can be estimated from simple measurements of muscle architecture such as muscle belly mass, fascicle length and physiological cross‐sectional area. While the hindlimb anatomy of the non‐human apes has been studied in some detail, a comparative study of the forelimb architecture across a number of species has never been undertaken. Here we present data from chimpanzees, bonobos, gorillas and an orangutan to ascertain if, and where, there are functional differences relating to their different locomotor repertoires and habitat usage. We employed a combination of analyses including allometric scaling and ancovas to explore the data, as the sample size was relatively small and heterogeneous (specimens of different sizes, ages and sex). Overall, subject to possible unidentified, confounding factors such as age effects, it appears that the non‐human great apes in this sample (the largest assembled to date) do not vary greatly across different muscle architecture parameters, even though they perform different locomotor behaviours at different frequencies. Therefore, it currently appears that the time spent performing a particular behaviour does not necessarily impose a dominating selective influence on the soft‐tissue portion of the musculoskeletal system; rather, the overall consistency of muscle architectural properties both between and within the Asian and African apes strengthens the case for the hypothesis of a possible ancient shared evolutionary origin for orthogrady under compressive and/or suspensory loading in the great apes.

Does footprint depth correlate with foot motion and pressure

Footprints are the most direct source of evidence about locomotor biomechanics in extinct vertebrates. One of the principal suppositions underpinning biomechanical inferences is that footprint geometry correlates with dynamic foot pressure, which, in turn, is linked with overall limb motion of the trackmaker. In this study, we perform the first quantitative test of this long-standing assumption, using topological statistical analysis of plantar pressures and experimental and computer-simulated footprints. In computer-simulated footprints, the relative distribution of depth differed from the distribution of both peak and pressure impulse in all simulations. Analysis of footprint samples with common loading inputs and similar depths reveals that only shallow footprints lack significant topological differences between depth and pressure distributions. Topological comparison of plantar pressures and experimental beach footprints demonstrates that geometry is highly dependent on overall print depth; deeper footprints are characterized by greater relative forefoot, and particularly toe, depth than shallow footprints. The highlighted difference between ‘shallow’ and ‘deep’ footprints clearly emphasizes the need to understand variation in foot mechanics across different degrees of substrate compliance. Overall, our results indicate that extreme caution is required when applying the ‘depth equals pressure’ paradigm to hominin footprints, and by extension, those of other extant and extinct tetrapods.

Regional peak plantar pressures are highly sensitive to region boundary definitions

Traditional pedobarographic analyses subsample pressure data over a number of discrete anatomical regions of interest (ROIs). To our knowledge, the sensitivity of these data to ROI boundary definitions has not been previously addressed. Eight subjects each performed 20 trials of self-paced walking; commercial software was used to define 10 ROIs for each of the 160 total peak pressure images, and regional peak pressures (RPPs) were extracted for each image (total: 1600 values). We then asked three specific questions regarding RPP sensitivity to ROI boundary definition: (1) Is the ROI centroid representative of the RPP location? (2) How frequently do RPPs lie on the ROI boundary? and (3) By how much do RPP values change if the ROI boundary is changed by one pixel (resolution: 5.08 x 7.62 mm)? We found that the RPP locations differed from the ROI centroid in 80% of the cases and that the RPPs lay on the ROI boundary with a probability of 65%. We also found that a single-pixel change in the ROI boundary caused a mean RPP change of 10.8%. The most sensitive region was the midfoot for which a single-pixel ROI change yielded a median 29.4% change in RPP. These results indicate that RPP data are biased by regionalization schemes, which delineate pressure fields based on anatomy rather than on the fields geometric properties, and ultimately that regionalization may constitute a poor method of quantifying complex pressure fields. RPP sensitivity should be considered when making statistical inferences regarding foot function.

Vector field statistics for objective center-of-pressure trajectory analysis during gait, with evidence of scalar sensitivity to small coordinate system rotations

Center of pressure (COP) trajectories summarize the complex mechanical interaction between the foot and a contacted surface. Each trajectory itself is also complex, comprising hundreds of instantaneous vectors over the duration of stance phase. To simplify statistical analysis often a small number of scalars are extracted from each COP trajectory. The purpose of this paper was to demonstrate how a more objective approach to COP analysis can avoid particular sensitivities of scalar extraction analysis. A previously published dataset describing the effects of walking speed on plantar pressure (PP) distributions was re-analyzed. After spatially and temporally normalizing the data, speed effects were assessed using a vector-field paired Hotellings T2 test. Results showed that, as walking speed increased, the COP moved increasingly posterior at heel contact, and increasingly laterally and anteriorly between ∼60 and 85% stance, in agreement with previous independent studies. Nevertheless, two extracted scalars disagreed with these results. Furthermore, sensitivity analysis found that a relatively small coordinate system rotation of 5.5° reversed the mediolateral null hypothesis rejection decision. Considering that the foot may adopt arbitrary postures in the horizontal plane, these sensitivity results suggest that non-negligible uncertainty may exist in mediolateral COP effects. As compared with COP scalar extraction, two key advantages of the vector-field approach are: (i) coordinate system independence, (ii) continuous statistical data reflecting the temporal extents of COP trajectory changes.