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The human calcaneus is robust and provides a prominent heel for effective bipedal locomotion, although the adjacent talus has no muscle attachments. However, there is incomplete information about the morphological changes in these prominent bones during embryo development. We examined serial histological sections of 23 human embryos and early-term fetuses (approximately 5-10 weeks' gestational age [GA]). At a GA of 5 weeks, the precartilage talus was parallel to and on the medial side of the calcaneus, which had a prolate spheroid shape and consisted of three masses. At a GA of 6 weeks, the cartilaginous talus extended along the proximodistal axis, and the tuber calcanei became long and bulky, with a small sustentaculum talus at the "distal" side. At a GA of 6 to 8 weeks, the sustentaculum had a medial extension below the talus so that the talus "rode over" the calcaneus. In contrast, the talus had a more complex shape, depending on the growth of adjacent bones. At a GA of 9 to 10 weeks, the talus was above the calcaneus, but the medial part still faced the plantar subcutaneous tissue because of the relatively small sustentaculum. Therefore, the final morphology appeared after an additional several weeks. Muscle activity seemed to facilitate growth of the tuber calcanei, but growth of the other parts of calcaneus, including the sustentaculum, seemed to depend on active proliferation at the different sites of cartilage. Multiple tendons and ligaments seemed to fix the talus so that it remained close to the calcaneus., (© 2024 Anatomical Society.)

The Kromdraai site in South Africa has yielded numerous early hominin fossils since 1938. As a part of recent excavations within Unit P, a largely complete early hominin calcaneus (KW 6302) was discovered. Due to its role in locomotion, the calcaneus has the potential to reveal important form/function relationships. Here, we describe KW 6302 and analyze its preserved morphology relative to human and nonhuman ape calcanei, as well as calcanei attributed to Australopithecus afarensis, Australopithecus africanus, Australopithecus sediba, Homo naledi, and the Omo calcaneus (either Paranthropus or early Homo). KW 6302 calcaneal morphology is assessed using numerous quantitative metrics including linear measures, calcaneal robusticity index, relative lateral plantar process position, Achilles tendon length reconstruction, and a three-dimensional geometric morphometric sliding semilandmark analysis. KW 6302 exhibits an overall calcaneal morphology that is intermediate between humans and nonhuman apes, although closer to modern humans. KW 6302 possesses many traits that indicate it was likely well-adapted for terrestrial bipedal locomotion, including a relatively flat posterior talar facet and a large lateral plantar process that is similarly positioned to modern humans. It also retains traits that indicate that climbing may have remained a part of its locomotor repertoire, such as a relatively gracile tuber and a large peroneal trochlea. Specimens from Kromdraai have been attributed to either Paranthropus robustus or early Homo; however, there are no definitively attributed calcanei for either genus, making it difficult to taxonomically assign this specimen. KW 6302 and the Omo calcaneus, however, fall outside the range of expected variation for an extant genus, indicating that if the Omo calcaneus was Paranthropus, then KW 6302 would likely be attributed to early Homo (or vice versa)., (© 2022 Anatomical Society.)

Bone structure dynamically adapts to its mechanical environment throughout ontogeny by altering the structure of trabecular bone, the three-dimensional mesh-like structure found underneath joint surfaces. Trabecular structure, then, can provide a record of variation in loading directions and magnitude; and in ontogenetic samples, it can potentially be used to track developmental shifts in limb posture. We aim to broaden the analysis of trabecular bone ontogeny by incorporating interactions between ontogenetic variation in locomotor repertoire, neuromuscular maturation, and life history. We examine the associations between these variables and age-related variation in trabecular structure in the calcaneus of Japanese macaques (Macaca fuscata). We used high-resolution micro-computed tomography scanning to image the calcaneus in a cross-sectional sample of 34 juvenile M. fuscata aged between 0 and 7 years old at the Primate Research Institute, Japan. We calculated whole bone averages of standard trabecular properties and generated whole-bone morphometric maps of bone volume fraction and Young's modulus. Trabecular structure becomes increasingly heterogeneous in older individuals. Bone volume fraction (BV/total volume [TV]) decreases during the first month of life and increases afterward, coinciding with the onset of independent locomotion in M. fuscata. At birth, primary Young's modulus is oriented orthogonal to the ossification center, but after locomotor onset bone structure becomes stiffest in the direction of joint surfaces and muscle attachments. Age-related variation in bone volume fraction is best predicted by an interaction between the estimated percentage of adult brain size, body mass, and locomotor onset. To explain our findings, we propose a model where interactions between age-related increases in body weight and maturation of the neuromuscular system alter the loading environment of the calcaneus, to which the internal trabecular structure dynamically adapts. This model cannot be directly tested based on our cross-sectional data. However, confirmation of the model by longitudinal experiments and in multiple species would show that trabecular structure can be used both to infer behavior from fossil morphology and serve as a valuable proxy for neuromuscular maturation and life history events like locomotor onset and the achievement of an adult-like gait. This approach could significantly expand our knowledge of the biology and behavior of fossil species., (© 2022 The Authors. Journal of Anatomy published by John Wiley & Sons Ltd on behalf of Anatomical Society.)

Quantifying morphological variation is critical for conducting anatomical research. Three-dimensional geometric morphometric (3D GM) landmark analyses quantify shape using homologous Cartesian coordinates (landmarks). Setting up a high-density landmark set and placing it on all specimens, however, can be a time-consuming task. Weighted spherical harmonics (SPHARM) provides an alternative method for analyzing the shape of such objects. Here we compare sliding semilandmark and SPHARM analyses of the calcaneus of Gorilla gorilla gorilla (n = 20), Pan troglodytes troglodytes (n = 20), and Homo sapiens (n = 20) to determine whether the SPHARM and sliding semilandmark analyses capture comparable levels of shape variation. We also compare both the sliding semilandmark and SPHARM analyses to a novel combination of the two methods, here termed SPHARM-sliding. In SPHARM-sliding, the vertices of the surface models produced from the SPHARM analysis (that are the same in number and relative location) are used as the starting landmark positions for a sliding semilandmark analysis. Calcaneal shape variation quantified by all three analyses was summarized using separate principal components analyses. Results were compared using the root mean square (RMS) and maximum distance between surface models of species averages scaled (up) to centroid size created from each analysis. The average RMS was 0.23 mm between sliding semilandmark and SPHARM average surface models, 0.19 mm between SPHARM and SPHARM sliding average surface models, and 0.22 mm between sliding semilandmark and SPHARM sliding average surface models. Although results indicate that all three analyses are comparable methods for 3D shape analysis, there are advantages and disadvantages to each. While the SPHARM analysis is less time-intensive, it is unable to capture the same level of detail around the sharp edges of articular facets on average surface models as the sliding semilandmark analysis. The SPHARM analysis also does not allow for individual articular facets to be analyzed in isolation. SPHARM-sliding, however, captures the same level of detail as the sliding semilandmark analysis, and (as in the sliding semilandmark analysis) allows for the evaluation of individual portions of bone. SPHARM is a comparable method to a 3D GM analysis for small, irregularly shaped bones, such as the calcaneus, and SPHARM-sliding allows for an expedited set up process for a sliding semilandmark analysis., (© 2021 Anatomical Society.)

The bones forming the talocrural joint (TCJ) and subtalar joint (STJ) are often assumed to be bilaterally symmetric. Therefore, the contralateral limb (i.e. the fibula, tibia, calcaneus and talus) is used as a template or an intra-subject control in clinical and research practice. However, the validity of the symmetry assumption is controversial, because insufficient information is available on the shape variations and bilateral (a)symmetry of the fibula, tibia, calcaneus and talus. Using three-dimensional spatially dense sampled representations of bone shapes extracted from bilateral computed tomography scans of 66 individuals (55 male, mean age: 61 ± 10 years; 11 female, mean age: 53 ± 15 years), we analyzed whether: (i) similar shape patterns exist in the left and right bones of the same type; (ii) gender has an effect on bone shape variations; (iii) intra-subject shape variation is smaller than that of inter-subject for a given shape variance direction. For the first set of analyses, all left and right instances of the same type of bone were considered as two separate groups, and statistically compared with each other on multiple aspects including group location (central tendency), variance-covariance scale (dispersion) and orientation (covariance structure) using distance-based permutational tests. For the second and third sets of analyses, all left and right bones of the same type were pooled into one group, and shape variations in the TCJ and STJ bones were extracted using principal component analysis. The effects of gender on age-adjusted bone shape differences were assessed using an analysis of covariance. Moreover, intra-class correlation was employed to evaluate intra- and inter-subject bone shape variations. For each bone type, both sides had similar shape patterns (P permutational -values > 0.05). After Bonferroni adjustment, gender led to shape differences, which were mainly in the lateral and medial condyles of the tibia (P = 0.003), the length and height of the calcaneus (P < 0.001), the posterior and anterior talar articular surfaces of the calcaneus (P = 0.001), and in the posterior aspect of the talus (P = 0.001). Intra-subject shape variations in the tibial tuberosity together with the diameter of the tibia, and the curvature of the fibula shaft and the diameter of the fibula were as high as those of inter-subject. This result suggests that the shape symmetry assumption could be violated for some specific shape variations in the fibula and tibia., (© 2018 The Authors. Journal of Anatomy published by John Wiley & Sons Ltd on behalf of Anatomical Society.)

Sheep and deer calcanei are finding increased use as models for studies of bone adaptation, including advancing understanding of how the strain (deformation) environment influences the ontogenetic emergence of biomechanically relevant structural and material variations in cortical and trabecular bone. These artiodactyl calcanei seem ideal for these analyses because they function like simply loaded short-cantilevered beams with net compression and tension strains on the dorsal and plantar cortices, respectively. However, this habitual strain distribution requires more rigorous validation because it has been shown by limited in vivo and ex vivo strain measurements obtained during controlled ambulation (typically walking and trotting). The conception that these calcanei are relatively simply and habitually loaded 'tension/compression bones' could be invalid if infrequent, though biologically relevant, loads substantially change the location of the neutral axis (NA) that separates 'compression' and 'tension' regions. The effect on calcaneus strains of the tension members (plantar ligament and flexor tendon) is also not well understood and measuring strains after transecting them could reveal that they significantly modulate the strain distribution. We tested the hypothesis that the NA location previously described during simulated on-axis loads of deer calcanei would exhibit limited variations even when load perturbations are unusual (e.g. off-axis loads) or extreme (e.g. after transection of the tension members). We also examined regional differences in the predominance of the three strain modes (tension, compression, and shear) in these various load conditions in dorsal, plantar, medial, and lateral cortices. In addition to considering principal strains (tension and compression) and maximum shear strains, we also considered material-axis (M-A) shear strains. M-A shear strains are those that are aligned along the long axis of the bone and are considered to have greater biomechanical relevance than maximum shear strains because failure theories of composite materials and bone are often based on stresses or strains in the principal material directions. We used the same load apparatus from our prior study of mule deer calcanei. Results showed that although the NA rotated up to 8° medially and 15° laterally during these off-axis loads, it did not shift dramatically until after transection of all tension members. When comparing results based on maximum shear strain data vs. M-A shear strain data, the dominant strain mode changed only in the plantar cortex - as expected (in accordance with our a priori view) it was tension when M-A shear strains were considered (shear : tension = 0.2) but changed to dominant shear when maximum shear strain data were considered (shear : tension = 1.3). This difference leads to different conclusions and speculations regarding which specific strain modes and magnitudes most strongly influence the emergence of the marked mineralization and histomorphological differences in the dorsal vs. plantar cortices. Consequently, our prior simplification of the deer calcaneus model as a simply loaded 'tension/compression bone' (i.e. plantar/dorsal) might be incorrect. In vivo and in finite element analyses are needed to determine whether describing it as a 'shear-tension/compression' bone is more accurate. Addressing this question will help to advance the artiodactyl calcaneus as an experimental model for bone adaptation studies., (Published 2018. This article is a U.S. Government work and is in the public domain in the USA.)

The purpose of this study was to quantify isolated coupling mechanisms of calcaneal adduction/abduction and calcaneal eversion/inversion to proximal bones in vitro. The in vitro approach is necessary because in vivo both movements appear together, making it impossible to determine the extent of their individual contribution to overall ankle joint coupling. Eight fresh frozen foot-leg specimens were tested. Data describing bone orientation and coupling mechanisms between segments were obtained using bone pin marker triads. The bone movement was described in a global coordinate system to examine the coupling between the calcaneus, talus and tibia. The strength of coupling was determined by means of the slope of a linear least squares fit to an angle-angle plot. The coupling coefficients in the present study indicate that not only calcaneal eversion/inversion (coupling coefficient: 0.68 ± 0.15) but to an even greater extent calcaneal adduction/abduction (coupling coefficient: 0.99 ± 0.10) was transferred into talus and tibial rotation, highlighting the relevance of calcaneal adduction for the overall ankle joint coupling. The results of this study present the possibility that controlling calcaneal adduction/abduction can affect talus and tibial rotation and therefore the possible genesis of overuse knee injuries., (© 2018 Anatomical Society.)

Findings on the twisting structure and insertional location of the AT on the calcaneal tuberosity are inconsistent. Therefore, to obtain a better understanding of the mechanisms underlying insertional Achilles tendinopathy, clarification of the anatomy of the twisting structure and location of the AT insertion onto the calcaneal tuberosity is important. The purpose of this study was to reveal the twisted structure of the AT and the location of its insertion onto the calcaneal tuberosity using Japanese cadavers. The study was conducted using 132 legs from 74 cadavers (mean age at death, 78.3 ± 11.1 years; 87 sides from men, 45 from women). Only soleus (Sol) attached to the deep layer of the calcaneal tuberosity was classified as least twist (Type I), both the lateral head of the gastrocnemius (LG) and Sol attached to the deep layer of the calcaneal tuberosity were classified as moderate twist (Type II), and only LG attached to the deep layer of the calcaneal tuberosity was classified as extreme twist (Type III). The Achilles tendon insertion onto the calcaneal tuberosity was classified as a superior, middle or inferior facet. Twist structure was Type I (least) in 31 legs (24%), Type II (moderate) in 87 legs (67%), and Type III (extreme) in 12 legs (9%). A comparison between males and females revealed that among men, 20 legs (24%) were Type I, 57 legs (67%) Type II, and eight legs (9%) Type III. Among women, 11 legs (24%) were Type I, 30 legs (67%) Type II, and four legs (9%) Type III. No significant differences were apparent between sexes. The fascicles of the Achilles tendon attach mainly in the middle facet. Anterior fibers of the Achilles tendon, where insertional Achilles tendinopathy is most likely, are Sol in Type I, LG and Sol in Type II, and LG only in Type III. This suggests the possibility that a different strain is produced in the anterior fibers of the Achilles tendon (calcaneal side) where insertional Achilles tendinopathy is most likely to occur in each type. We look forward to elucidating the mechanisms generating insertional Achilles tendinopathy in future biomedical studies based on the present results., (© 2016 Anatomical Society.)

The osteogenic relationship between the lateral plantar process and the peroneal tubercle has been an uncertainty for researchers over several decades. While some argue there to be no developmental relationship between these two calcaneal structures, others have suggested that there is an inverse relationship, the lateral plantar process forming from a part of the peroneal tubercle. However, no previous studies have offered quantitative measurements to test these hypotheses. In this study, we measured the size of the peroneal tubercle, retrotrochlear eminence, and the size and area of the lateral plantar process in 73 subjects using magnetic resonance imaging (MRI). Navicular height was measured using weight-bearing radiographs as a measurement of longitudinal arch in 35 of these subjects. Age, body mass, and body mass index (BMI) were also recorded for all subjects. We determined that there was a significant positive correlation between the lateral plantar process and size of the peroneal tubercle, body mass, and BMI. Thus, assertions that there is an inverse relationship between the size of the lateral plantar process and the peroneal tubercle are here unfounded. We also determined there to be a positive correlation between the peroneal tubercle and both the size of the retrotrochlear eminence and the height of the navicular. In conclusion, we relate these novel findings to hominin fossil calcanei and discuss the evolutionary and biomechanical implications., (© 2013 Anatomical Society.)

Six mechanically significant skeletal variables were measured on the calcanei from 60 Felidae specimens (22 species) to determine whether these variables were scaled to body mass, and to assess whether differential scaling exists. The power equation (y = a · x(b) ) was used to analyse the scaling of the six variables to body mass; we compared traditional regression methods (standardised major axis) to phylogenetically independent contrasts. In agreement with previous studies that compared these methodologies, we found no significant differences between methods in the allometric coefficients (b) obtained. Overall, the scaling pattern of the felid calcaneus conformed to the predictions of the geometric similarity hypothesis, but not entirely to those of the elastic similarity hypothesis. We found that the moment arm of the ankle extensors scaled to body mass with an exponent not significantly different from 0.40. This indicated that the tuber calcanei scaled to body mass faster than calcaneus total length. This explained why the effective mechanical advantage of the ankle extensors increased with body mass, despite the fact that limb posture does not change in felid species. Furthermore, this finding was consistent with the hypothesis of the isometric scaling of ground reaction forces. No evidence for differential scaling was found in any of the variables studied. We propose that this reflected the similar locomotor pattern of all felid species. Thus, our results suggested that the differences in allometric coefficients for 'large' and 'small' mammals were in fact caused by different types of locomotion among the species included in each category., (© 2012 The Authors. Journal of Anatomy © 2012 Anatomical Society.)

Calcanei of mature mule deer have the largest mineral content (percent ash) difference between their dorsal 'compression' and plantar 'tension' cortices of any bone that has been studied. The opposing trabecular tracts, which are contiguous with the cortices, might also show important mineral content differences and microscopic mineralization heterogeneity (reflecting increased hemi-osteonal renewal) that optimize mechanical behaviors in tension vs. compression. Support for these hypotheses could reveal a largely unrecognized capacity for phenotypic plasticity - the adaptability of trabecular bone material as a means for differentially enhancing mechanical properties for local strain environments produced by habitual bending. Fifteen skeletally mature and 15 immature deer calcanei were cut transversely into two segments (40% and 50% shaft length), and cores were removed to determine mineral (ash) content from 'tension' and 'compression' trabecular tracts and their adjacent cortices. Seven bones/group were analyzed for differences between tracts in: first, microscopic trabecular bone packets and mineralization heterogeneity (backscattered electron imaging, BSE); and second, trabecular architecture (micro-computed tomography). Among the eight architectural characteristics evaluated [including bone volume fraction (BVF) and structural model index (SMI)]: first, only the 'tension' tract of immature bones showed significantly greater BVF and more negative SMI (i.e. increased honeycomb morphology) than the 'compression' tract of immature bones; and second, the 'compression' tracts of both groups showed significantly greater structural order/alignment than the corresponding 'tension' tracts. Although mineralization heterogeneity differed between the tracts in only the immature group, in both groups the mineral content derived from BSE images was significantly greater (P < 0.01), and bulk mineral (ash) content tended to be greater in the 'compression' tracts (immature 3.6%, P = 0.03; mature 3.1%, P = 0.09). These differences are much less than the approximately 8% greater mineral content of their 'compression' cortices (P < 0.001). Published data, suggesting that these small mineralization differences are not mechanically important in the context of conventional tests, support the probability that architectural modifications primarily adapt the tracts for local demands. However, greater hemi-osteonal packets in the tension trabecular tract of only the mature bones (P = 0.006) might have an important role, and possible synergism with mineralization and/or microarchitecture, in differential toughening at the trabeculum level for tension vs. compression strains., (© Published 2012. This article is a US Government work and is in the public domain in the USA. Journal of Anatomy © 2012 Anatomical Society.)

Natural loading of the calcanei of deer, elk, sheep and horses produces marked regional differences in prevalent/predominant strain modes: compression in the dorsal cortex, shear in medial-lateral cortices, and tension/shear in the plantar cortex. This consistent non-uniform strain distribution is useful for investigating mechanisms that mediate the development of the remarkable regional material variations of these bones (e.g. collagen orientation, mineralization, remodeling rates and secondary osteon morphotypes, size and population density). Regional differences in strain-mode-specific microdamage prevalence and/or morphology might evoke and sustain the remodeling that produces this material heterogeneity in accordance with local strain characteristics. Adult calcanei from 11 animals of each species (deer, elk, sheep and horses) were transversely sectioned and examined using light and confocal microscopy. With light microscopy, 20 linear microcracks were identified (deer: 10; elk: six; horse: four; sheep: none), and with confocal microscopy substantially more microdamage with typically non-linear morphology was identified (deer: 45; elk: 24; horse: 15; sheep: none). No clear regional patterns of strain-mode-specific microdamage were found in the three species with microdamage. In these species, the highest overall concentrations occurred in the plantar cortex. This might reflect increased susceptibility of microdamage in habitual tension/shear. Absence of detectable microdamage in sheep calcanei may represent the (presumably) relatively greater physical activity of deer, elk and horses. Absence of differences in microdamage prevalence/morphology between dorsal, medial and lateral cortices of these bones, and the general absence of spatial patterns of strain-mode-specific microdamage, might reflect the prior emergence of non-uniform osteon-mediated adaptations that reduce deleterious concentrations of microdamage by the adult stage of bone development., (© Published 2011. This article is a U.S. Government work and is in the public domain in the USA. Journal of Anatomy © 2011 Anatomical Society of Great Britain and Ireland.)

The source of pain and the background to the pain mechanisms associated with mid-portion Achilles tendinopathy have not yet been clarified. Intratendinous degenerative changes are most often addressed when present. However, it is questionable if degeneration of the tendon itself is the main cause of pain. Pain is often most prominent on the medial side, 2-7cm from the insertion onto the calcaneus. The medial location of the pain has been explained to be caused by enhanced stress on the calcaneal tendon due to hyperpronation. However, on this medial side the plantaris tendon is also located. It has been postulated that the plantaris tendon might play a role in these medially located symptoms. To our knowledge, the exact anatomy and relationship between the plantaris- and calcaneal tendon at the level of complaints have not been anatomically assessed. This was the purpose of our study. One-hundred and seven lower extremities were dissected. After opening the superficial fascia and paratendon, the plantaris tendon was bluntly released from the calcaneal tendon moving distally. The incidence of the plantaris tendon, its course, site of insertion and possible connections were documented. When with manual force the plantaris tendon could not be released, it was defined as a 'connection' with the calcaneal tendon. In all specimens a plantaris tendon was identified. Nine different sites of insertion were found, mostly medial and fan-shaped onto the calcaneus. In 11 specimens (10%) firm connections were found at the level of the calcaneal tendon mid-portion. Clinical and histological studies are needed to confirm the role of the plantaris tendon in mid-portion Achilles tendinopathy., (© 2011 The Authors. Journal of Anatomy © 2011 Anatomical Society of Great Britain and Ireland.)

The attachment of the Achilles tendon is part of an 'enthesis organ' that reduces stress concentration at the hard-soft tissue interface. The organ also includes opposing sesamoid and periosteal fibrocartilages, a bursa and Kager's fat pad. In addition, the deep crural and plantar fasciae contribute to Achilles stress dissipation and could also be regarded as components. Here we describe the sequence in which these various tissues differentiate. Serial sections of feet from spontaneously aborted foetuses (crown rump lengths 22-322 mm) were examined. All slides formed part of an existing collection of histologically sectioned embryological material, obtained under Spanish law and housed in the Universidad Complutense, Madrid. From the earliest stages, it was evident that the Achilles tendon and plantar fascia had a mutual attachment to the calcaneal perichondrium. The first components of the enthesis organ to appear (in the 45-mm foetus) were the retrocalcaneal bursa and the crural fascia. The former developed by cavitation within the mesenchyme that later gave rise to Kager's fat pad. The tip of the putative fat pad protruded into the developing bursa in the 110-mm foetus and fully differentiated adipocytes were apparent in the 17-mm foetus. All three fibrocartilages were first recognisable in the 332-mm foetus--at which time adipogenesis had commenced in the heel fat pad. The sequence in which the various elements became apparent suggests that bursal formation and the appearance of the crural fascia may be necessary to facilitate the foot movements that subsequently lead to fibrocartilage differentiation. The later commencement of adipogenesis in the heel than in Kager's pad probably reflects the non-weight environment in utero. The direct continuity between plantar fascia and Achilles tendon that is characteristic of the adult reflects the initial attachment of both structures to the calcaneal perichondrium rather than to the skeletal anlagen itself.

The distribution of type II collagen in sagittal sections of the Achilles tendon has been used to reconstruct the three-dimensional (3D) shape and position of three fibrocartilages (sesamoid, periosteal and enthesis) associated with its insertion. The results showed that there is a close correspondence between the shape and position of the sesamoid and periosteal fibrocartilages--probably because of their functional interdependence. The former protects the tendon from compression during dorsiflexion of the foot, and the latter protects the superior tuberosity of the calcaneus. When the zone of calcified enthesis fibrocartilage and the subchondral bone are mapped in 3D, the reconstructions show that there is a complex pattern of interlocking between pieces of calcified fibrocartilage and bone at the insertion site. We suggest that this is of fundamental importance in anchoring the tendon to the bone, because the manner in which a tendon insertion develops makes it unlikely that many collagen fibres pass across the tissue boundary from tendon to bone. When force is transmitted to the bone from a loaded tendon, it is directed towards the plantar fascia by a series of highly orientated trabeculae that are clearly visible in 3D in thick resin sections.

Sarcomere number and sarcomere length were studied in six groups of kittens ranging in age from 10 minutes to 5 months and compared with those of adult cats. Although the soleus muscle is a slow contracting muscle and the tibialis anterior a fast contracting muscle, both have previously been shown to have the same range relative to ankle movement. For a given angle of articulation the sarcomere length was found to be the same at all ages except perhaps for the newly born. In contrast, the sarcomere number differed considerably, being much higher in the older animals. The relationship between active tension and muscle length was also measured, and again no difference was found between the muscles at any age, although the shape of the curves for the soleus and tibialis anterior was different.

The internal structure of the tarsal bones has been studied to investigate their cancellous architecture. It is revealed that these bones have fine and coarse meshworks and even a tendency for obliteration of the trabecular pattern in the bones lying distal to this midtarsal joint. Internal structure of the talus does not show an arched pattern of bony lamellae. An increased density of bony lamellae in the internal structure of the navicular bone could result from excessive stress, enforced by its close relationship to the three cuneiform bones.

The variations in the talar articular facets on 401 normal Indian calcanei have been observed and classified. Type 1 calcanei have one continuous facet on the sustentaculum tali extending on the antero-medial corner of the distal part of the calcaneus (268 cases--67%). Type 2 calcanei have two articular facets, one on the sustentaculum tali, and another on the distal part of the calcaneus (104 cases--26%). Type 3 calcanei have a single well defined facet on the sustentaculum tali (21 cases--5%). Type 4 calcanei have a single facet on the sustenaculum tali extending on to the antero-medial corner of the distal part of the calcaneus, but this facet is continuous with the posterior talar facet of the calcaneus (8 cases--2%).

The preferential orientation of the apatite crystals in the lower tibia, talus and calcaneus, as determined by neutron diffraction, serves as an indicator of the lines of stress in the foot. The main stress flows down from the tibia to the heel and the ball of the foot and there is noteworthy orientation along the line of the plantar aponeurosis which acts as a tie. Orientation is particularly marked above the point of attachment of the Achilles tendon, where the edge of the calcaneus functions as a beam. The centre of the talus serves as a junctional region for forces and is relatively unoriented.

The absolute density of the right and left os calcis has been measured in a group of healthy, sedentary volunteers. There was a significant difference between the density of the right and left os calcis in 90% of the subjects. Statistically, when all of the subjects were considered, the side on which the os calcis was denser was also the side of hand preference. Since there was no evidence, in the present study, of the environment having encouraged the use of one foot rather than the other, it is concluded that the concordance between upper limb dominance and greater density of the ipsilateral os calcis must have been determined before birth.

The fossil record supports the conclusions derived from the study of extant species that the Primates evolved a unique suite of characters in the articulations of the foot. The tarsal bones of African Miocene apes show specializations characteristic of hominoid evolution and provide reasonable precursors for the morphology of Pan, Gorilla and even Pongo. The OH8 foot is essentially ape-like in it major features, with many close resemblances to Pan. Although fairly clearly from a bipedal primate, it lacked important functional specializations found in the human foot.

It is shown that in form and function the articular complexes of the monotreme foot are pre-adaptive to the therian condition, but the echidna differs by having a tuber calcaneus which is directed downward and distally. The cynodont foot (TR. 8) and that of the Triassic mammal Eozostrodon seem to possess the essential articular features present in monotremes, but they are assembled rather differently. In both, tuber calcaneus was apparently directed downwards. It follows that monotremes were probably derived from some way along the lineage usually, but inappropriately, termed "Theria'. A calcaneofibular articulation is present in kangaroos, certain shrews, elephant shrews, rabbits and artiodactyls. In all of them it is an apomorphic condition involving annexation of part of the posterior talar facet on the calcaneus by the fibula, which invariably shows some degree of amalgamation with the tibia. It is shown that the trochlear process of the mammalian calcaneus has the dual function of providing origin for the m. flexor accessorius and acting as a supporting shelf for the bundle of peroneal tendons. It is almost certainly a derivative of the lateral flange on the cynodont calcaneus, which presumably had a comparable function. In man, the process is fragmented, one of its derivatives being the lateral process of the calcaneal tuber which shows varying degrees of migration towards the medial process and amalgamation with it. The importance of these morphological features is discussed in relation to their use in cladistic analysis and their relevance to theories of the early evolution of the mammals.