title: Biomechanical implications of mineral content and microstructural variations in cortical bone of horse, elk, and sheep calcanei.
abstract: Artiodactyl and perissodactyl calcanei have been recently introduced as models for examining bone for mechanically mediated adaptation. We have reported substantial regional variations in cortical bone microstructure and mineral content within the same cross-section of mule deer calcanei. In part, these variations may be adaptations accommodating the customary presence of predominantly tension, compression, and shear strain modes in mutually exclusive cortical locations. Calcanei from skeletally mature horses, elk, and sheep were examined in order to corroborate these previous findings.
  
  
  
  From each species, one calcaneus was obtained from each of 13 animals. Each bone was cut transversely near mid-shaft into two segments and examined for mineral (ash) content. From each species, an additional segment obtained from each of 7 of the original 13 bones was examined for microstructure using 50x backscattered electron images. Regions examined included the compression (cranial), tension (caudal), and medial and lateral (shear) cortices. Periosteal (P), middle (M), and endosteal (E) regions were also examined separately within the compression and tension cortices. Quantified microstructural parameters included: (1) secondary osteon population density (OPD), (2) fractional area of secondary bone (FASB), (3) porosity, (4) population density of new remodeling events (NRE = resorption spaces and newly forming secondary osteons), and (5) secondary osteon diameter and minimum-to-maximum chord ratio.
  
  
  
  Results in each species showed variations that are considered to be mechanically important and are similar to those reported in mule deer calcanei. Mineral content data suggest that remodeling activity in the compression, medial, and lateral cortices was occurring at a slower rate than remodeling in the tension cortex. In comparison to the tension cortices, the compression cortices have approximately 6.0% higher mineral content (P < 0.007) and 35% higher OPD (P < 0.01). Additionally, the compression cortices have more nearly perfectly round osteons and lower FASB, porosity, NRE, and osteon diameter (P < 0.05; except for FASB in horse where P = 0.087 and NRE in sheep where P = 0.520). However, patterns of microstructural variations between intracortical regions (P, M, E) are inconsistent when compared to data reported in mule deer calcanei. Microstructural characteristics between the medial and lateral cortices were similar although some significant differences were identified. In general, the microstructure of the medial and lateral cortices differ from the neighboring compression and tension cortices.
  
  
  
  Differences in mineral content and microstructure between opposing compression and tension cortices of these three species resemble differences previously reported in mule deer calcanei. The majority of the microstructural variations can be explained in the context of strain-magnitude-based rules of Frost's Mechanostat Theory of mechanically induced bone adaptation. These variations may also be strongly influenced by the strain mode predominating in each cortical location. The hypothesis that intracortical material adaptations are correlated with progressive transcortical strain magnitude variations is not supported by the inconsistent transcortical variations in material organization. These interpretations do not preclude the possibility that other specific strain features may contribute to a complex adaptive signal.
authors.affiliation: Bone and Joint Research Laboratory, VA Medical Center, Salt Lake City, Utah 84148, USA.
authors.forename: J G
authors.initials: JG
authors.lastname: Skedros
authors.forename: S C
authors.initials: SC
authors.lastname: Su
authors.forename: R D
authors.initials: RD
authors.lastname: Bloebaum
mesh_terms: D000818:Animals; D001696:Biomechanical Phenomena; D016723:Bone Remodeling / Q000502:physiology; D002111:Calcaneus / Q000033:anatomy & histology* / Q000378:metabolism* / Q000502:physiology; D002113:Calcification, Physiologic / Q000502:physiology*; D003670:Deer / Q000033:anatomy & histology* / Q000502:physiology; D005260:Female; D006736:Horses / Q000033:anatomy & histology* / Q000502:physiology; D008297:Male; D012756:Sheep / Q000033:anatomy & histology* / Q000502:physiology