Regional maps of rib cortical bone thickness and cross‐sectional geometry

Here we present detailed regional bone thickness and cross‐sectional measurements from full adult ribs using high resolution CT scans processed with a cortical bone mapping technique. Sixth ribs from 33 subjects ranging from 24 to 99 years of age were used to produce average cortical bone thickness maps and to provide average ± 1SD corridors for expected cross‐section properties (cross‐sectional areas and inertial moments) as a function of rib length. Results obtained from CT data were validated at specific rib locations using direct measurements from cut sections. Individual thickness measurements from CT had an accuracy (mean error) and precision (SD error) of ?0.013 ± 0.167 mm (R² coefficient of determination of 0.84). CT‐based measurement errors for rib cross‐sectional geometry were ?0.1 ± 13.1% (cortical bone cross‐sectional area) and 4.7 ± 1.8% (total cross‐sectional area). Rib cortical bone thickness maps show the expected regional variation across a typical rib's surface. The local mid‐rib maxima in cortical thickness along the pleural rib aspect ranged from range 0.9 to 2.6 mm across the study population with an average map maximum of 1.4 mm. Along the cutaneous aspect, rib cortical bone thickness ranged from 0.7 to 1.9 mm with an average map thickness of 0.9 mm. Average cross‐sectional properties show a steady reduction in total cortical bone area from 10% along the rib's length through to the sternal end, whereas overall cross‐sectional area remains relatively constant along the majority of the rib's length before rising steeply towards the sternal end. On average, male ribs contained more cortical bone within a given cross‐section than was seen for female ribs. Importantly, however, this difference was driven by male ribs having larger overall cross‐sectional areas, rather than by sex differences in the bone thickness observed at specific local cortex sites. The cortical bone thickness results here can be used directly to improve the accuracy of current human body and rib models. Furthermore, the measurement corridors obtained from adult subjects across a wide age range can be used to validate future measurements from more widely available image sources such as clinical CT where gold standard reference measures (e.g. such as direct measurements obtained from cut sections) are otherwise unobtainable.     

颞骨的断面解剖学研究及其临床意义

目的：进一步明确颞骨的断面形态和各结构的解剖关系，为影像诊断和手术治疗提供解剖学依据。方法：将6例成人头颅标本制备成层厚0．5mm的耳颞部火棉胶横断切片，逐一观测。结果：以HRCT为参考，选取11个典型层面：①上半规管层面；②上鼓室层面；③外半规管层面；④锤砧关节层面；⑤内耳道底下部层面；⑥前庭窗层面；⑦匙突层面；⑧后鼓室层面；⑨蜗窗层面；⑩鼓膜脐层面；(11)咽鼓管层面。对层面内主要结构的断面形态及其毗邻关系进行了描述。结论：颞骨火棉胶薄切片能很好地显示颞骨的解剖结构及其毗邻关系，对该区的影像诊断和手术治疗有重要的参考价值。     

Strain-mode-specific mechanical testing and the interpretation of bone adaptation in the deer calcaneus

The artiodactyl (deer and sheep) calcaneus is a model that helps in understanding how many bones achieve anatomical optimization and functional adaptation. We consider how the dorsal and plantar cortices of these bones are optimized in quasi-isolation (the conventional view) versus in the context of load sharing along the calcaneal shaft by “tension members” (the plantar ligament and superficial digital flexor tendon). This load-sharing concept replaces the conventional view, as we have argued in a recent publication that employs an advanced analytical model of habitual loading and fracture risk factors of the deer calcaneus. Like deer and sheep calcanei, many mammalian limb bones also experience prevalent bending, which seems problematic because the bone is weaker and less fatigue-resistant in tension than compression. To understand how bones adapt to bending loads and counteract deleterious consequences of tension, it is important to examine both strain-mode-specific (S-M-S) testing (compression testing of bone habitually loaded in compression; tension testing of bone habitually loaded in tension) and non-S-M-S testing. Mechanical testing was performed on individually machined specimens from the dorsal “compression cortex” and plantar “tension cortex” of adult deer calcanei and were independently tested to failure in one of these two strain modes. We hypothesized that the mechanical properties of each cortex region would be optimized for its habitual strain mode when these regions are considered independently. Consistent with this hypothesis, energy absorption parameters were approximately three times greater in S-M-S compression testing in the dorsal/compression cortex when compared to non-S-M-S tension testing of the dorsal cortex. However, inconsistent with this hypothesis, S-M-S tension testing of the plantar/tension cortex did not show greater energy absorption compared to non-S-M-S compression testing of the plantar cortex. When compared to the dorsal cortex, the plantar cortex only had a higher elastic modulus (in S-M-S testing of both regions). Therefore, the greater strength and capacity for energy absorption of the dorsal cortex might “protect” the weaker plantar cortex during functional loading. However, this conventional interpretation (i.e., considering adaptation of each cortex in isolation) is rejected when critically considering the load-sharing influences of the ligament and tendon that course along the plantar cortex. This important finding/interpretation has general implications for a better understanding of how other similarly loaded bones achieve anatomical optimization and functional adaptation.     

Global deletion of Panx3 produces multiple phenotypic effects in mouse humeri and femora

Pannexins form single‐membrane channels that allow passage of small molecules between the intracellular and extracellular compartments. Of the three pannexin family members, Pannexin3 (Panx3) is the least studied but is highly expressed in skeletal tissues and is thought to play a role in the regulation of chondrocyte and osteoblast proliferation and differentiation. The purpose of our study is to closely examine the in vivo effects of Panx3 ablation on long bone morphology using micro‐computed tomography. Using Panx3 knockout (KO) and wildtype (WT) adult mice, we measured and compared aspects of phenotypic shape, bone mineral density (BMD), cross‐sectional geometric properties of right femora and humeri, and lean mass. We found that KO mice have absolutely and relatively shorter diaphyseal shafts compared with WT mice, and relatively larger areas of muscle attachment sites. No differences in BMD or lean mass were found between WT and KO mice. Interestingly, KO mice had more robust femora and humeri compared with WT mice when assessed in cross‐section at the midshaft. Our results clearly show that Panx3 ablation produces phenotypic effects in mouse femora and humeri, and support the premise that Panx3 has a role in regulating long bone growth and development.     

Collagen Fiber Orientation in Primate Long Bones

Studies of variation in orientation of collagen fibers within bone have lead to the proposition that these are preferentially aligned to accommodate different kinds of load, with tension best resisted by fibers aligned longitudinally relative to the load, and compression best resisted by transversely aligned fibers. However, previous studies have often neglected to consider the effect of developmental processes, including constraints on collagen fiber orientation (CFO), particularly in primary bone. Here we use circularly polarized light microscopy to examine patterns of CFO in cross‐sections from the midshaft femur, humerus, tibia, radius, and ulna in a range of living primate taxa with varied body sizes, phylogenetic relationships and positional behaviors. We find that a preponderance of longitudinally oriented collagen is characteristic of both periosteal primary and intracortically remodeled bone. Where variation does occur among groups, it is not simply understood via interpretations of mechanical loads, although prioritized adaptations to tension and/or shear are considered. While there is some suggestion that CFO may correlate with body size, this relationship is neither consistent nor easily explicable through consideration of size‐related changes in mechanical adaptation. The results of our study indicate that there is no clear relationship between CFO and phylogenetic status. One of the principle factors accounting for the range of variation that does exist is primary tissue type, where slower depositing bone is more likely to comprise a larger proportion of oblique to transverse collagen fibers. Anat Rec, 300:1189–1207, 2017. © 2017 Wiley Periodicals, Inc.     

Relationships among microstructural properties of bone at the human midshaft femur

Mineralization density and collagen fibre orientation are two aspects of a bone's microstructural organization that influence its mechanical properties. Previous studies by our group have demonstrated a distinctly non-random, though highly variable, spatial distribution of these two variables in the human femoral cortex. In this study of 37 specimens, these variables are examined relative to one another in order to determine whether regions of bone demonstrating higher or lower mineralization density also demonstrate a prevalence of either transversely or longitudinally oriented collagen fibres. An analysis of rank-transformed collagen fibre orientation (as determined by circularly polarized light) and mineralization density (as determined by backscattered electron microscopy) data sets demonstrated that areas of low mineralization density (predominantly in the anterior-lateral cortex) tended to correspond to regions of higher proportions of longitudinally oriented collagen fibres. Conversely, areas of higher mineralization density (postero-medially) tended to correspond to regions of higher proportions of transversely oriented collagen fibres. High variability in the sample led to generally low correlations between the two data sets, however. A second analysis focused only on the orientation of collagen fibres within poorly mineralized bone (representing bone that was newly formed). This analysis demonstrated a lower proportion of transverse collagen fibres in newly formed bone with age, along with some significant regional differences in the prevalence of collagen fibres of either orientation. Again high variability characterized the sample. These results are discussed relative to the hypothesized forces experienced at the midshaft femur.     

Determination of the orientation of collagen fibers in human bone

A human calcaneus bone, consisting of hydroxyapatite and collagen fibers, was successively sliced into samples in a direction perpendicular to the long axis of the bone and parallel to the long axis of the human lower limb. The transmitted microwave intensities of 12 GHz, reflecting the dielectric property, were measured for the slice samples using Osaki's microwave method (Tappi J., 1987 ; 70:105–108). The complex dielectric constant of 12 GHz of the collagen fiber film was much greater than that of hydroxyapatite disc, which demonstrated that the dielectric anisotropy observed for the sliced bone was mainly affected by the collagen fibers. The angular dependence of the transmitted microwave intensity gives the orientation angle reflecting the direction of the collagen‐fiber orientation, and the degree of orientation reflecting the anisotropic property of collagen fibers. The orientation angle and the degree of orientation for the slice samples changed with changing position along the long axis of the calcaneus bone. The direction of orientation deviated to the lateral side at the heel part of the left calcaneus, and to the medial side at the middle part. The degree of orientation is relatively high at the heel part and low at the middle. Such results give a two‐dimensional distribution of collagen‐fiber orientation in the left calcaneus, and suggest that the direction and degree of orientation are closely related to the direction and magnitude of the stress applied to the bone, respectively. Anat Rec 266:103–107, 2002. © 2002 Wiley‐Liss, Inc.     

Three‐dimensional reconstruction of Haversian systems in human cortical bone using synchrotron radiation‐based micro‐CT: morphology and quantification of branching and transverse connections across age

This study uses synchrotron radiation‐based micro‐computed tomography (CT) scans to reconstruct three‐dimensional networks of Haversian systems in human cortical bone in order to observe and analyse interconnectivity of Haversian systems and the development of total Haversian networks across different ages. A better knowledge of how Haversian systems interact with each other is essential to improve understanding of remodeling mechanisms and bone maintenance; however, previous methodological approaches (e.g. serial sections) did not reveal enough detail to follow the specific morphology of Haversian branching, for example. Accordingly, the aim of the present study was to identify the morphological diversity of branching patterns and transverse connections, and to understand how they change with age. Two types of branching morphologies were identified: lateral branching, resulting in small osteon branches bifurcating off of larger Haversian canals; and dichotomous branching, the formation of two new osteonal branches from one. The reconstructions in this study also suggest that Haversian systems frequently target previously existing systems as a path for their course, resulting in a cross‐sectional morphology frequently referred to as ‘type II osteons’. Transverse connections were diverse in their course from linear to oblique to curvy. Quantitative assessment of age‐related trends indicates that while in younger human individuals transverse connections were most common, in older individuals more evidence of connections resulting from Haversian systems growing inside previously existing systems was found. Despite these changes in morphological characteristics, a relatively constant degree of overall interconnectivity is maintained throughout life. Altogether, the present study reveals important details about Haversian systems and their relation to each other that can be used towards a better understanding of cortical bone remodeling as well as a more accurate interpretation of morphological variants of osteons in cross‐sectional microscopy. Permitting visibility of reversal lines, synchrotron radiation‐based micro‐CT is a valuable tool for the reconstruction of Haversian systems, and future analyses have the potential to further improve understanding of various important aspects of bone growth, maintenance and health.     

Ontogenetic structural and material variations in ovine calcanei: A model for interpreting bone adaptation

Experimental models are needed for resolving relative influences of genetic, epigenetic, and nonheritable functionally induced (extragenetic) factors in the emergence of developmental adaptations in limb bones of larger mammals. We examined regional/ontogenetic morphologic variations in sheep calcanei, which exhibit marked heterogeneity in structural and material organization by skeletal maturity. Cross‐sections and lateral radiographs of an ontogenetic series of domesticated sheep calcanei (fetal to adult) were examined for variations in biomechanically important structural (cortical thickness and trabecular architecture) and material (percent ash and predominant collagen fiber orientation) characteristics. Results showed delayed development of variations in cortical thickness and collagen fiber orientation, which correlate with extragenetic factors, including compression/tension strains of habitual bending in respective dorsal/plantar cortices and load‐related thresholds for modeling/remodeling activities. In contrast, the appearance of trabecular arches in utero suggests strong genetic/epigenetic influences. These stark spatial/temporal variations in sheep calcanei provide a compelling model for investigating causal mechanisms that mediate this construction. In view of these findings, it is also suggested that the conventional distinction between genetic and epigenetic factors in limb bone development be expanded into three categories: genetic, epigenetic, and extragenetic factors. Anat Rec, 2007. © 2007 Wiley‐Liss, Inc.     

Bone shaft bending strength index is unaffected by exercise and unloading in mice

Anthropologists frequently use the shaft bending strength index to infer the physical activity levels of humans living in the past from their lower limb bone remains. This index is typically calculated as the ratio of bone shaft second moments of area about orthogonal principal axes (i.e. I_max/I_min). Individuals with high I_max/I_min values are inferred to have been very active, whereas individuals with low values are inferred to have been more sedentary. However, there is little direct evidence that activity has a causal and predictable effect on the shaft bending strength index. Here, we report the results of two experiments that were designed to test the model within which anthropologists commonly interpret the shaft bending strength index. In the first experiment, mice were treated daily with treadmill exercise for 1 month to simulate a high‐activity lifestyle. In the second experiment, in an attempt to simulate a low‐activity lifestyle, functional weight‐bearing was removed from the hindlimbs of mice for 1 month. Femoral mid‐shaft structure was determined with μCT. We found that while exercise resulted in significant enhancement of I_max and I_min compared with controls, it failed to significantly increase the I_max/I_min index. Similarly, stunted bone growth caused by unloading resulted in significantly diminished I_max and I_min compared with controls, but low activity did not lead to significantly decreased I_max/I_min compared with normal activity. Together, these results suggest that caution is required when the bone shaft bending strength index is used to reconstruct the activity levels of past humans.     

The effects of immobilization on vascular canal orientation in rat cortical bone

It is well established that bone is capable of adapting to changes in loading; however, little is known regarding how loading specifically affects the internal 3D microarchitecture of cortical bone. The aim of this study was to experimentally test the hypothesis that loading is a determinant of the 3D orientation of primary vascular canals in the rat tibial diaphysis. Left tibiae from 10 rats (30 weeks old) that had been immobilized (sciatic neurectomy) for 27 weeks, right SHAM-operated tibiae from these same rats (internal control) and right tibiae from 10 normal age-matched rats (external control) were scanned by micro-CT. Mean canal orientation (for the whole bone segment and by region), percent porosity, canal diameter and canal separation were quantitatively assessed in 3D. Canal orientation in the immobilized tibiae was significantly (P < 0.001) more radial (by 9.9°) compared to the external controls but did not differ from the internal controls (P = 0.310). Comparing the external and internal controls, orientation was significantly (P < 0.05) more radial in the internal control group (by 6.8°). No differences were found for percent porosity and canal separation. Canal diameter was significantly greater in the immobilized vs. internal (P < 0.001) and external control (P < 0.001) tibiae. The differences in orientation relative to the external controls indicated that the organization of cortical bone in the rat is affected by loading. Although the predicted difference in canal orientation was not detected between immobilized and internal control groups, the distributions of individual canal orientations, from which the mean values were derived, revealed distinctive patterns for all three groups. The internal controls exhibited an intermediate position between the immobilized and external controls, suggesting that paralysis on the contralateral side resulted in altered loading relative to the normal state represented by the external control. This was also evident in a regional analysis by quadrant. The loaded bones had the same cross-sectional shape; however, their internal structure differed. These results provide novel insights into the impact of loading on the 3D organization of primary cortical bone and have implications for understanding the relation between cortical bone adaptation, disease and mechanical properties.     

Exploring Femoral Diaphyseal Shape Variation in Wild and Captive Chimpanzees by Means of Morphometric Mapping: A Test of Wolff's Law

Long bone shafts (diaphyses) serve as load‐bearing structures during locomotion, implying a close relationship between diaphyseal form and its locomotor function. Diaphyseal form‐function relationships, however, are complex, as they are mediated by various factors such as developmental programs, evolutionary adaptation, and functional adaptation through bone remodeling during an individual's lifetime. The effects of the latter process (“Wolff's Law”) are best assessed by comparing diaphyseal morphologies of conspecific individuals under different locomotor regimes. Here we use morphometric mapping (MM) to analyze the morphology of entire femoral diaphyses in an ontogenetic series of wild and captive common chimpanzees (Pan troglodytes troglodytes). MM reveals patterns of variation of diaphyseal structural and functional properties, which cannot be recognized with conventional cross‐sectional analysis and/or geometric morphometric methods. Our data show that diaphyseal shape, cortical bone distribution and inferred cross‐sectional biomechanical properties vary both along ontogenetic trajectories and independent of ontogeny. Mean ontogenetic trajectories of wild and captive chimpanzees, however, were found to be statistically identical. This indicates that the basic developmental program of the diaphysis is not altered by different loading conditions. Significant differences in diaphyseal shape between groups could only be identified in the distal diaphysis, where wild chimpanzees exhibit higher mediolateral relative to anteroposterior cortical bone thickness. Overall, thus, the hypothesis that Wolff's Law predominantly governs long bone diaphyseal morphology is rejected. Anat Rec, 2011. © 2011 Wiley‐Liss, Inc.     

胶原变性对骨的变形与断裂的影响

骨中有机成分特别是胶原对其生物力学性质具有重要影响。骨中胶原的物理化学和生物学性质的变化均会导致骨的变形和断裂的危险性的增加。然而,胶原的影响往往易被忽视。本文综述了近年来胶原变性对骨的变形和断裂行为影响的研究进展。分析了胶原变性的一些主要影响因素(如年龄、激素、辐射、病变和药物等),进一步分析了由于胶原的变性而引起的胶原纤维排列方向的改变、多肽链的异构化、三级螺旋结构的破坏以及交联物含量的减少等因素最终导致骨变形与断裂的微观机制。     

Ontogenetic relationships between in vivo strain environment, bone histomorphometry and growth in the goat radius

Vertebrate long bone form, at both the gross and the microstructural level, is the result of many interrelated influences. One factor that is considered to have a significant effect on bone form is the mechanical environment experienced by the bone during growth. The work presented here examines the possible relationships between in vivo bone strains, bone geometry and histomorphology in the radii of three age/size groups of domestic goats. In vivo bone strain data were collected from the radii of galloping goats, and the regional cortical distribution of peak axial strain magnitudes, radial and circumferential strain gradients, and longitudinal strain rates related to regional patterns in cortical growth, porosity, remodelling and collagen fibre orientation. Although porosity and remodelling decreased and increased with age, respectively, these features showed no significant regional differences and did not correspond to regional patterns in the mechanical environment. Thicker regions of the radius's cortex were significantly related to high strain levels and higher rates of periosteal, but not endosteal, growth. However, cortical growth and strain environment were not significantly related. Collagen fibre orientation varied regionally, with a higher percentage of transverse fibres in the caudal region of the radius and primarily longitudinal fibres elsewhere, and, although consistent through growth, also did not generally correspond to regional strain patterns. Although strain magnitudes increased during ontogeny and regional strain patterns were variable over the course of a stride, mean regional strain patterns were generally consistent with growth, suggesting that regional growth patterns and histomorphology, in combination with external loads, may play some role in producing a relatively 'predictable' strain environment within the radius. It is further hypothesized that the absence of correlation between regional histomorphometric patterns and the measured strain environments is the result of the variable mechanical environment. However, the potential effects of other physiological and mechanical factors, such as skeletal metabolism and adjacent muscle insertions, that can influence the gross and microstructural morphology of the radius during ontogeny, cannot be ignored.     

Ontogenetic changes of geometrical and mechanical characteristics of the avian femur: a comparison between precocial and altricial birds

The mechanical performance of limb bones is closely associated with an animal's locomotor capability and is thus important to our understanding of animal behaviour. This study combined a geometrical analysis and three‐point bending tests to address the question of how the mechanical performance of the femurs of Japanese quail (Coturnix coturnix japonica) and pigeon (Columba livia domestica) respond to changing functional demands during ontogeny. Results showed that hatchling quails had stiff bone tissues, and the femoral ultimate loads scaled negatively with body mass, corresponding to high functional demands during early growth. The hatchling pigeon femora had weak material properties but they showed a dramatic increase in Young's modulus during growth. Consequently, although femoral cross‐sectional geometry showed negative allometry, the ultimate loads scaled positively with body mass. Older pigeons had more circular bone cross‐sections than younger pigeons, probably due to load stimulation changes occurred shortly after the onset of locomotion. Negative allometry and isometry of the cross‐sectional geometry of hind limb bones were observed in flying birds and ground‐dwelling birds, respectively. The correspondence between geometrical change and locomotor pattern suggests that ontogenetic changes in cross‐sectional geometry may be an effective indicator of avian locomotor behaviour.     

Cross‐relaxation parameters in cortical bone assessed with different MR sequences

This study aimed to show evidence of MR cross‐relaxation effects in cortical bone and to compare different MR sequences for the quantification of cross‐relaxation parameters. Measurements were performed on bovine diaphysis samples with spectroscopic methods (inversion‐recovery, off‐resonance saturation) and with a variable flip angle (VFA) UTE imaging method on a 4.7 T laboratory‐assembled scanner. Cross‐relaxation parameter assessment was carried out via a two‐pool model simulation with a matrix algebra approach. A proton signal amplitude of 28 Mol/L was observed (equivalent water fraction of 25%). It was attributed to collagen‐bound water, with values of ~ 0.3 ms, a “long‐T₂” proton pool, in exchange with protons from the collagen macromolecules ( of 10–20 μs). Magnetization transfer (MT) effects were detected with all sequences. The best precision of model parameters was obtained with off‐resonance saturation; the fraction of collagen methylene protons was found in the range of 22–28% and the transverse relaxation time for collagen methylene protons was 11 μs (1% precision). The model parameters obtained were compatible with VFA‐UTE results but could not be assessed with acceptable accuracy and precision using this method. In vivo MT quantification using off‐resonance saturation with a single B₁ amplitude and offset frequency may provide information about the relative amount of collagen per unit volume in cortical bone.     

Secondary osteon variants and remodeling in human bone

Histomorphometric analysis of human cortical bone has documented the occurrence of secondary osteon variants. These include drifting osteons which form tails as they move erratically through the cortex and Type II osteons which show partial resorption and redeposition within the cement line of the osteon. Little is known about the biological significance of these variants. Prior studies suggested correlations with age, biomechanics, diet, and mineral homeostasis. No study has yet tested for osteon variant associations with static measures of bone remodeling. In this study, thin sections (n = 112) of the posterior femur representing a late English Medieval adult human osteological collection, subdivided by age, sex, and socio-economic status, were examined to determine whether remodeling indicators reconstructed from osteon parameters (area, diameter, area ratios) and densities differed between categories of presence or absence of Type II and drifting osteon variants. Of the 112 sections, 33 presented with Type II osteons, and 38 had drifting osteons. Sporadic statistically significant results were identified. Haversian canal:osteon area ratio differed (p = 0.017) with Type II osteon presence, Type II osteons were more prevalent in males than females (p = 0.048), and drifting osteons were associated with smaller osteon (p = 0.049) and Haversian canal area (p = 0.05). These results may be explained through some biological (sex) and social (status) processes such as a period of physiological recovery (e.g., following lactation, malnutrition). However, the general lack of consistent relationships between osteon variants and remodeling indicators suggests they occur as a result of natural variation.     

Histological organization and its relationship to function in the femur of Alligator mississippiensis

Histological analysis of a growth series of alligator femora tests the correlation between strain milieu and microstructure. From mid‐diaphyseal cross‐sections of these femora (n = 7), vascular canal orientation and density as well as collagen fibre organization were recorded. Throughout ontogeny, the proportion of transverse–spiral (TS) collagen in the dorsal cortex is significantly greater than it is in the ventral cortex (P = 0.008). This regional difference in the proportion of TS collagen is correlated with a regional difference in the state of peak principal strain (compressive or tensile). Nevertheless, the predominant orientation of collagen fibres is longitudinal, which is inconsistent with biomechanical hypotheses that involve peak principal or shear strains. Although the density and orientation of vascular canals do not show significant regional differences (P = 0.26 and P = 0.26, respectively), as with collagen orientation, the vascular canal orientation is predominantly longitudinal. The longitudinal organization of both the vascular canals and the collagen fibres is probably a consequence of longitudinal shifting of subperiosteal osteoid during femoral lengthening. When taken together, these data suggest that growth dynamics is the dominant influence on the histological organization of primary bony tissues in alligator femora.     

Ontogeny of hallucal metatarsal rigidity and shape in the rhesus monkey (Macaca mulatta) and chimpanzee (Pan troglodytes)

Life history variables including the timing of locomotor independence, along with changes in preferred locomotor behaviors and substrate use during development, influence how primates use their feet throughout ontogeny. Changes in foot function during development, in particular the nature of how the hallux is used in grasping, can lead to different structural changes in foot bones. To test this hypothesis, metatarsal midshaft rigidity [estimated from the polar second moment of area (J) scaled to bone length] and cross‐sectional shape (calculated from the ratio of maximum and minimum second moments of area, I_max/I_min) were examined in a cross‐sectional ontogenetic sample of rhesus macaques (Macaca mulatta; n = 73) and common chimpanzees (Pan troglodytes; n = 79). Results show the hallucal metatarsal (Mt1) is relatively more rigid (with higher scaled J‐values) in younger chimpanzees and macaques, with significant decreases in relative rigidity in both taxa until the age of achieving locomotor independence. Within each age group, Mt1 rigidity is always significantly higher in chimpanzees than macaques. When compared with the lateral metatarsals (Mt2–5), the Mt1 is relatively more rigid in both taxa and across all ages; however, this difference is significantly greater in chimpanzees. Length and J scale with negative allometry in all metatarsals and in both species (except the Mt2 of chimpanzees, which scales with positive allometry). Only in macaques does Mt1 midshaft shape significantly change across ontogeny, with older individuals having more elliptical cross‐sections. Different patterns of development in metatarsal diaphyseal rigidity and shape likely reflect the different ways in which the foot, and in particular the hallux, functions across ontogeny in apes and monkeys.