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 materialaxis (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 cortexas 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.Journal of ...
Customary nonuniform distributions of physiological bone strains are thought to evoke heterogeneous material adaptation in diaphyseal cortices of some limb bones. Recent studies of artiodactyl calcanei have suggested that the regional prevalence of specific mechanical strain features such as mode and magnitude correlate with specific variations in cortical bone ultrastructure, microstructure and mineralization. These data are also consistent with predictions of current algorithms of mechanically induced bone adaptation. However, detailed characterization of the customary functional strain environment of these bones is needed to understand better the mechanisms of these adaptations. An in vitro loading method and rosette strain gauges were used to record principal strains, maximum shear strains and principal strain angles at multiple locations on ten calcanei of adult male mule deer (Odocoileus hemionus hemionus). Each hind limb was fixed in an apparatus to mimic the mid-support phase of the gait and loaded via the Achilles tendon over a broad range of functional loads (0 to 2943 N). Strains were recorded on the craniolateral, craniomedial, caudal, medial and lateral cortices at mid-diaphysis. Loading variations included the progressive elimination of the ligament and tendon along the caudal calcaneus. The results showed that the cranial cortex experiences longitudinal compressive strains that are nearly equal to the principal minimum strains and that the caudal cortex receives longitudinal tensile strains that are nearly equal to the principal maximum strains. With a 981 N load, the mean principal compressive strain on the cranial cortex was −636+/−344 micro(ε) (mean +/− s.d., N=9) and the mean principal tensile strain on the caudal cortex was 1112+/−68 micro;(ε)x (N=9). In contrast to the cranial and caudal cortices, principal strains in the medial and lateral cortices displayed relatively large deviations from the longitudinal axis (medial, 24 degrees cranial; lateral, 27 degrees caudal). Although shear strains predominated at all gauge sites, variations in maximum shear strains showed no apparent regional pattern or consistent regional predominance. The plantar ligament and tendon of the superficial digital flexor muscle were shown to have important load-sharing functions. These results demonstrate that the functionally loaded artiodactyl calcaneus generally behaves like a cantilevered beam with longitudinal compression and tension strains predominating in opposing cranial and caudal cortices, respectively. Differences in osteon remodeling rates, osteon morphology and mineral content reported previously between the cranial and caudal cortices correlate, in part, with the magnitudes of the principal compressive and tensile strains, respectively. However, material differences that distinguish the medial and lateral cortices from the cranial and caudal cortices could not be primarily attributed to locally increased shear strains as previously suggested. Variations in osteon and/or collagen fiber orientation may correlate more strongly with principal strain direction.
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