The split-line phenomenon and the microscopic structure of bone.
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Living adult Eskimos from St. Lawrence Island, North Alaska, and Canada undergo an earlier and more rapid rate of age-related bone mineral loss compared to U.S. whites. Further, it has been shown that Eskimos and Indians differ in patterns of osteon remodeling at the Haversian envelope. Femoral bone cores from adult Eskimos skeletons from St. Lawrence Island (n = 53), Kodiak Island (n = 92), Baffin Island (n = 44), and Southampton Island (n = 69) were analyzed and the results compared with those obtained from cores from U.S. whites (n = 144). Cortical thickness, bone mineral content of cores, cortical bone density, secondary osteon and Haversian canal number and area were quantified for each core. Ages at death were estimated by histological methods and compared with the ages at death estimated by morphological methods for the Eskimo skeletons. Known ages at death were compared with histologically estimated ages at death for the U.S. white series. St. Lawrence Island and Kodiak Island (Yupik speakers) Eskimo cortical thickness values were significantly (P less than .05) greater than Baffin Island and Southampton Island (Inupiaq speakers) Eskimos cortical thickness values but less than the cortical thickness values for U.S. whites. The bone mineral content of the Southampton Eskimos femoral cores was the lowest found in this study. Histological analysis of the femoral bone sections showed that Eskimos contain more osteons per unit area than U.S. whites. No differences in osteon size were noted between the two populations. Differences in patterns of osteon remodeling between Eskimos and whites were inferred. Age at death estimation by histological methods in Eskimos using U.S. white regression equations yielded age estimates in poor agreement with those obtained by morphological methods.
Mechanical factors are implicated in the control of osteonal (Haversian) remodeling. Twenty primate species, representing a diverse range of locomotor behaviors, have been examined to determine whether the arrangement of secondary osteonal bone at femoral mid-diaphysis reflects behaviorally related biomechanical influences. The proportion of bone cortex occupied by secondary osteons (percent osteonal bone) is related to the biomechanical environment of these femora. Percent osteonal bone delineates groups which correspond to other skeletal and kinesiological categorizations of primate locomotion and distinguishes among arboreal quadrupedal, terrestrial quadrupedal, suspensory, and bipedal primates.
Ahlqvist and Damsten's (1969) modification of the Kerley (1965) method for histological age estimation uses percent osteonal bone, rather than actual osteon counts, in order to eliminate the difficulty of distinguishing between intact and fragmentary osteons. Since their method has been developed for the femur only, and several more recent methods have been proposed that utilize percent osteonal bone, a study was undertaken to ascertain the relative value of percent osteonal bone compared with osteon counts to estimate age at death for the radius, tibia, and fibula. First the question of how much of the cross-section of a bone should be sampled was addressed by comparing the results of regression against age for percent osteonal bone derived from sampling only four fields with those derived from the entire cross-section of the radius. A significant age association was found only when the entire cross-section was sampled. In order to evaluate the relative merit of using either percent osteonal bone, or osteon counts to estimate age, each variable was regressed against age. Significant correlation coefficients were found for all three bones when the independent variable was osteon counts. When percent osteonal bone was employed, a significant correlation was found only for the radius. Stepwise linear regression found osteon counts for the fibula alone to be the best age predictor. Finally, a repeated measures analysis of variance revealed that percent osteonal bone and osteon counts both differ among the three bones within an individual. Based upon these results, osteon counts, rather than percent osteonal bone, should be the variable of choice when developing histological age predicting methods.
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Variation in the size of structures within mature cortical bone is relevant to our understanding of apparent differences between human samples, and it is relevant to the development of histologically based age-estimation methods. It was proposed that variation may reflect effects of physical activity, through biomechanical and/or metabolic mechanisms. If these factors are local, femoral osteon area (On.Ar) should be more histologically variable than On.Ar in ribs. Ribs should show a higher variation in Haversian canal area (H.Ar) if they are sites of more remodeling activity and hence of arrested refilling of secondary osteons at time of death. This study compares On.Ar and H.Ar of secondary osteons from femora (15) and ribs (29) from 44 Holocene (Later Stone Age) foragers from South Africa (M = 19, F = 25) to values from paired femora and ribs from historic samples (Spitalfields and St. Thomas, 20 pairs from each). Fixed-effects analysis of variance demonstrates rib On.Ar to be significantly smaller than femur, but with no sex or age effects. The femur-to-rib On.Ar ratio is lower for the Holocene foragers than for the two modern samples because of relatively large rib On.Ar. Femora and ribs from the same skeleton normally show femoral On.Ar larger than rib On.Ar (37/44 pairs). Mean femoral values of On.Ar are more diverse than rib On.Ar values, but within-sample coefficients of variation are similar. Values for H.Ar are highly variable and do not reflect anatomical site, age, sex, or population effects. The patterning of osteon size does not appear to be linked to physical activity or to different rates of metabolic activity within the skeleton, at least not in a straightforward way.
Histological methods for the estimation of age at death using cortical bone are based on the evaluation of microstructural changes over time. Since histological analysis is a destructive method, most techniques attempt to limit the amount of cortical bone needed for analysis. Sample location, however, can have a significant effect on the accuracy of these methods. Furthermore, research demonstrates that both intersection and intrasection variation is present at the midshaft of the femur, which is the primary location for estimating age at death in humans. This research determines the extent of regional variation within the adult human femur and its effect on age estimation. Secondary osteon lamellae and Haversian canal ratio and cortical thickness were quantified. Thompson's All Males Left Femur regression equation was used to estimate age. Results show that significant regional variation occurs in the estimated ages derived from the posterior aspect of the femoral shaft and significant intrasection variation occurs in age estimates from the mid and mid-distal cross-sections. Thus, the inter and intrasection variation that occurs in bone remodeling within the femoral cortex has the potential to produce significant differences amongst age estimates taken from various femoral diaphyseal locations compared to the age estimated from the standard location used in Thompson's core method (1978). The results indicate that the use of this histological method is dependant on the ability to correctly identify the four anatomical locations, but the extracted core used for age estimation is not necessarily confined to the anterior midshaft.
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A procedure for differential staining of decalcified bone with silver nitrate showed major histological features which appeared to correspond closely to microradiographic images. The extent to which this is actually the case was investigated directly by preparing microradiographs of ground sections of baboon and dog radii and then decalcifying and staining the same sections. The many detailed similarities indicate that this staining procedure is a useful adjunct to microradiography. Thus, poorly mineralized osteons or layers of circumferential lamellae are darker stained by silver nitrate, and the variably mineralized layers of circumferential lamellae are closely duplicated by light and dark bands in the stained sections. These similarities imply that there is a relationship between the mineral density of bone and some condition of the organic matrix which is probably related to maturation changes in the collagen.
This paper compares patterns of histologic change in an archaic Native American population with those in modern white populations. Histologic sections were removed from core biopsies taken from the anterior femoral cortex of an archeologic sample of Pecos Indians. The data demonstrate many microstructural similarities between the Pecos and modern populations, even though they were genetically and culturally distinct. Pecos women had small Haversian canals and large osteon mean wall thickness, with no clear evidence of an intracortical bone volume deficit even in the older age groups, although significant marrow cavity expansion occurred in both males and females with age. No striking relationships were found between bone tissue changes and gross geometric changes with age. The data suggest that a more active life-style is associated with greater osteon mean wall thickness or osteon population density, but that it alone does not protect against significant bone loss on the cortical-endosteal surface.
There is no detailed information available concerning the variations in bone, the Haversian canal, and osteocyte populations in different-sized osteons. In this study a total of 398 secondary osteons were measured in archived rib sections from nine white men (20-25 years old). The sections were stained with basic fuchsin. The parameters included the osteon area (On.Ar), Haversian canal area (HC.Ar) and perimeter (HC.Pm), bone area (B.Ar), and osteocyte lacunar number (Lc.N). From these primary measurements the following indices were deduced: 1) lacunar number per bone area (Lc.N/B.Ar) and per osteon (Lc.N/On); 2) the ratio between Haversian canal perimeter and bone area (HC.Pm/B.Ar); and 3) the fraction of Haversian canal area (HC.Ar/On.Ar) and its complement, the fraction of bone area (B.Ar/On.Ar). The results showed that the osteons varied greatly in size, but very little in the fraction of bone area. Regression analyses showed that HC.Ar, HC.Pm, and Lc.N/On were positively associated with On.Ar (P < 0.001 for all). A significant negative correlation was found between On.Ar and Lc.N/B.Ar (P < 0.05) and HC.Pm/B.Ar (P < 0.0001). HC.Ar and HC.Pm increased significantly with increasing Lc.N/On (both P < 0.0001) rather than Lc.N/B.Ar. Lc.N/B.Ar had a significant positive correlation with HC.Ar/On.Ar (P < 0.05) and HC.Pm/B.Ar (P < 0.01). We conclude that: 1) the size of the osteon is determined by the quantum of bone removed by osteoclasts, 2) the osteon is well designed for molecular exchange, and 3) a well designed osteon may be produced via the regulation of bone apposition by osteocytes during the process of osteon refilling.
This study examines relationships between bone morphology and mechanically mediated strain/fluid-flow patterns in an avian species. Using mid-diaphyseal transverse sections of domestic turkey ulnae (from 11 subadults and 11 adults), we quantified developmental changes in predominant collagen fiber orientation (CFO), mineral content (%ash), and microstructure in cortical octants or quadrants (i.e., %ash). Geometric parameters were examined using whole mid-diaphyseal cross-sections. The ulna undergoes habitual bending and torsion, and demonstrates nonuniform matrix fluid-flow patterns, and high circumferential strain gradients along the neutral axis (cranial-caudal) region at mid-diaphysis. The current results showed significant porosity differences: 1) greater osteocyte lacuna densities (N.Lac/Ar) (i.e., "non-vascular porosity") in the caudal and cranial cortices in both groups, 2) greater N.Lac/Ar in the pericortex vs. endocortex in mature bones, and 3) greater nonlacunar porosity (i.e., "vascular porosity") in the endocortex vs. pericortex in mature bones. Vascular and nonvascular porosities were not correlated. There were no secondary osteons in subadults. In adults, the highest secondary osteon population densities and lowest %ash occurred in the ventral-caudal, caudal, and cranial cortices, where shear strains, circumferential strain gradients, and fluid displacements are highest. Changes in thickness of the caudal cortex explained the largest proportion of the age-related increase in cranial-caudal breadth; the thickness of other cortices (dorsal, ventral, and cranial) exhibited smaller changes. Only subadult bones exhibited CFO patterns corresponding to habitual tension (ventral) and compression (dorsal). These CFO variations may be adaptations for differential mechanical requirements in "strain-mode-specific" loading. The more uniform oblique-to-transverse CFO patterns in adult bones may represent adaptations for shear strains produced by torsional loading, which is presumably more prevalent in adults. The micro- and ultrastructural heterogeneities may influence strain and fluid-flow dynamics, which are considered proximate signals in bone adaptation.