Osbjorn M. Pearson

Optimization of bone growth and remodeling in response to loading in tapered mammalian limbs

SUMMARY How bones respond dynamically to mechanical loading through changes in shape and structure is poorly understood, particularly with respect to variations between bones. Structurally, cortical bones adapt in vivo to their mechanical environments primarily by modulating two processes, modeling and Haversian remodeling. Modeling, defined here as the addition of new bone, may occur in response to mechanical stimuli by altering bone shape or size through growth. Haversian remodeling is thought to be an adaptation to repair microcracks or prevent microcrack propagation. Here, we examine whether cortical bone in sheep limbs modulates periosteal modeling and Haversian remodeling to optimize strength relative to mass in hind-limb midshafts in response to moderate levels of exercise at different growth stages. Histomorphometry was used to compare rates of periosteal growth and Haversian remodeling in exercised and sedentary treatment groups of juvenile, subadult and young adult sheep. In vivo strain data were also collected for the tibia and metatarsal midshafts of juvenile sheep. The results suggest that limb bones initially optimize responses to loading according to the varying power requirements associated with adding mass at different locations. In juveniles, exercise induces higher rates of periosteal modeling in proximal midshafts and higher rates of Haversian remodeling in distal midshafts. Consequently, distal element midshafts experience higher strains and, presumably, have lower safety factors. As animals age, periosteal modeling rates decline and Haversian remodeling rates increase, but moderate levels of mechanical loading stimulate neither process significantly.

Bone density studies in zooarchaeology

Abstract Accurate bone density data are essential for assessing the influence of destructive processes in archaeological faunal assemblages. Unfortunately, the diversity of methods employed by different researchers to derive density values has resulted in recent confusion. Two recent publications in this journal [J Archaeol Sci 29 (2002) 883; J Archaeol Sci 29 (2002) 979] exemplify this state of misunderstanding. Both studies argued that the role of bone density in shaping archaeological faunal assemblages has been largely overrated, but both based their conclusions on density values that were inaccurately derived. The former employed a method of calculating bone density that has been largely discredited over the past decade [J Archaeol Sci 29 (2002) 883]. Within a larger discussion of zooarchaeological methodology, the latter provided an assessment of the current state of bone density research that inappropriately characterized the discrepancies between available sets of density data as a reflection of the differences between two technologies—photon densitometry and computed tomography [J Archaeol Sci 29 (2002) 979]. The actual dichotomy exists—irrespective of the technology employed—between studies that account for variation in the shape of bone cross-sections and those that do not. The different sets of density data currently available to zooarchaeologists vary tremendously in their accuracy. We review and evaluate the different techniques employed in the research of bone density patterns of mammalian fauna. Computed tomography produces the most accurate density data. For elements without medullary cavities, photon densitometry may provide density values of similar accuracy but only if a method of cross-sectional shape-adjustment is applied.

Postcranial remains and the origin of modern humans

The nature, timing, and location of the origin of modern humans has been the subject of intense controversy for the last 15 years.1–4 Genetic data and new radiometric dates for key fossils that lie beyond the range of radiocarbon dating have substantially added to the knowledge derived from the fossil evidence documenting the transition from archaic to modern humans. These new data, however, have failed to resolve the problem in its entirety. Most authorities now accept that Africa played an important, and probably central, role in the origin of modern humans.7–13 The genetic evidence seems to be particularly emphatic that an African population that existed between 200,000 and 100,000 years ago (100 ka) is ancestral to all living humans.6,7 Controversy still surrounds the question of how much, if at all, archaic humans from outside of Africa, such as Neandertals, late archaic Chinese hominins such as Jinniushan, and the Indonesian Ngandong hominins, may have contributed to the morphological and genetic diversity present in living populations and the morphology of the earliest fossils of modern humans.10

A description of the Omo I postcranial skeleton, including newly discovered fossils

Recent fieldwork in the Kibish Formation has expanded our knowledge of the geological, archaeological, and faunal context of the Omo I skeleton, the earliest known anatomically modern human. In the course of this fieldwork, several additional fragments of the skeleton were recovered: a middle manual phalanx, a distal manual phalanx, a right talus, a large and a small fragment of the left os coxae, a portion of the distal diaphysis of the right femur that conjoins with the distal epiphysis recovered in 1967, and a costal fragment. Some researchers have described the original postcranial fragments of Omo I as anatomically modern but have noted that a variety of aspects of the specimens morphology depart from the usual anatomy of many recent populations. Reanalysis confirms this conclusion. Some of the unusual features in Omo I--a medially facing radial tuberosity, a laterally flaring facet on the talus for the lateral malleolus, and reduced dorsovolar curvature of the base of metacarpal I--are shared with Neandertals, some early modern humans from Skhul and Qafzeh, and some individuals from the European Gravettian, raising the possibility that Eurasian early modern humans inherited these features from an African predecessor rather than Neandertals. The fragment of the os coxae does not unambiguously diagnose Omo Is sex: the greater sciatic notch is intermediate in form, the acetabulum is large (male?), and a preauricular sulcus is present (female?). The preserved portion of the left humerus suggests that Omo I was quite tall, perhaps 178-182 cm, but the first metatarsal suggests a shorter stature of 162-173 cm. The morphology of the auricular surface of the os coxae suggests a young adult age.

The importance of accounting for the area of the medullary cavity in cross-sectional geometry: A test based on the femoral midshaft.

In cross-sectional geometric (CSG) studies, both the subperiosteal and endosteal contours are considered important factors in determining bone bending rigidity. Recently, regression equations predicting CSG properties from a sections external dimensions were developed in a world-wide sample of human long bones. The results showed high correlations between some subperiosteally derived and actual CSG parameters. We present a theoretical model that further explores the influence of endosteal dimensions on CSG properties. We compare two hypothetical femoral midshaft samples with the same total subperiosteal area but with percentages of cortical bone at the opposite ends of published human variation for population sample means. Even in this relatively uncommon scenario, the difference between the samples in the resultant means for predicted femoral polar second moment of area (J) appears to be modest: power analysis indicates that a minimum sample size of 61 is needed to detect the difference 90% of the time via a t-test. Moreover, endosteal area can be predicted--although with substantial error--from periosteal area. Despite this error, including this relationship in subperiosteally derived estimates of J produces sample mean estimates close to true mean values. Power analyses reveal that when similar samples are used to develop prediction equations, a minimum sample of hundreds or more may be needed to distinguish a predicted mean J from the true mean J. These results further justify the use of regression equations estimating J from periosteal contours when analyzing behaviorally induced changes in bone rigidity in ancient populations, when it is not possible to measure endosteal dimensions. However, in other situations involving comparisons of individual values, growth trends, and senescence, where relative cortical thickness may vary greatly, inclusion of endosteal dimensions is still important.

Brains versus brawn: An empirical test of Barker's brain sparing model

The Barker model of the in utero origins of diminished muscle mass in those born small invokes the adaptive “sparing” of brain tissue development at the expense of muscle. Though compelling, to date this model has not been directly tested. This article develops an allometric framework for testing the principal prediction of the Barker model—that among those born small muscle mass is sacrificed to spare brain growth—then evaluates this hypothesis using data from the third National Health and Nutrition Examination Survey (NHANES III). The results indicate clear support for a negative relationship between the allometric development of the two tissues; however, a further consideration of conserved mammalian fetal circulatory patterns suggests the possibility that system‐constrained patterns of developmental damage and “bet‐hedging” responses in affected tissues may provide a more adequate explanation of the results. Far from signaling the end of studies of adaptive developmental programming, this perspective may open a promising new avenue of inquiry within the fields of human biology and the developmental origins of health and disease. Am. J. Hum. Biol., 2010.

Exercise-Induced Bone Formation Is Poorly Linked to Local Strain Magnitude in the Sheep Tibia

Functional interpretations of limb bone structure frequently assume that diaphyses adjust their shape by adding bone primarily across the plane in which they are habitually loaded in order to minimize loading-induced strains. Here, to test this hypothesis, we characterize the in vivo strain environment of the sheep tibial midshaft during treadmill exercise and examine whether this activity promotes bone formation disproportionately in the direction of loading in diaphyseal regions that experience the highest strains. It is shown that during treadmill exercise, sheep tibiae were bent in an anteroposterior direction, generating maximal tensile and compressive strains on the anterior and posterior shaft surfaces, respectively. Exercise led to significantly increased periosteal bone formation; however, rather than being biased toward areas of maximal strains across the anteroposterior axis, exercise-related osteogenesis occurred primarily around the medial half of the shaft circumference, in both high and low strain regions. Overall, the results of this study demonstrate that loading-induced bone growth is not closely linked to local strain magnitude in every instance. Therefore, caution is necessary when bone shaft shape is used to infer functional loading history in the absence of in vivo data on how bones are loaded and how they actually respond to loading.

Are human hands and feet affected by climate? A test of Allen's rule

OBJECTIVES In recent years, several studies have shown that populations from cold, high-latitude regions tend to have relatively shorter limbs than populations from tropical regions, with most of the difference due to the relative length of the zeugopods (i.e., radius, ulna, tibia, fibula). This pattern has been explained either as the consequence of long-term climatic selection or of phenotypic plasticity, with temperature having a direct effect on bone growth during development. The aims of this study were to test whether this pattern of intra-limb proportions extended to the bones of the hands and feet, and to determine whether the pattern remained significant after taking into account the effects of neutral evolutionary processes related to population history. MATERIALS AND METHODS Measurements of the limb bones, including the first metatarsal and metacarpal, were collected for 393 individuals from 10 globally distributed human populations. The relationship between intra-limb indices and minimum temperature was tested using generalized least squares regression, correcting for spatial autocorrelation. RESULTS The results confirmed previous observations of a temperature-related gradient in intra-limb proportions, even accounting for population history. This pattern extends to the hands, with populations from cold regions displaying a relatively shorter and stockier first metacarpal; however, the first metatarsal appears to be wider but not shorter in cold-adapted populations. DISCUSSION The results suggest that climatic adaptation played a role in shaping variation in limb proportions between human populations. The different patterns shown by the hands and feet might be due to the presence of evolutionary constraints on the foot to maintain efficient bipedal locomotion.

Developmental plasticity in fat patterning of Ache children in response to variation in interbirth intervals: a preliminary test of the roles of external environment and maternal reproductive strategies.

A firm link between small size at birth and later more centralized fat patterning has been established in previous research. Relationships between shortened interbirth intervals and small size at birth suggest that maternal energetic prioritization may be an important, but unexplored determinant of offspring fat patterning. Potential adaptive advantages to centralized fat storage (Baker et al., 2008 : In: Trevathan W, McKenna J, Smith EO, editors. Evolutionary Medicine and Health: New Perspectives. New York: Oxford) suggest that relationships with interbirth intervals may reflect adaptive responses to variation in patterns of maternal reproductive effort. Kuzawa ( 2005 : Am J Hum Biol 17:5–21; 2008 : In: Trevathan W, McKenna J, Smith EO, editors. Evolutionary Medicine and Health: New Perspectives. New York: Oxford) has argued that maternal mediation of the energetic quality of the environment is a necessary component of developmental plasticity models invoking predictive adaptive responses (Gluckman and Hanson 2004 : Trends Endocrinol Metab 15:183–187). This study tested the general hypothesis that shortened interbirth intervals would predict more centralized fat patterning in offspring. If long‐term maternally mediated signals are important determinants of offspring responses, then we expected to observe a relationship between the average interbirth interval of mothers and offspring adiposity, with no relationship with the preceding interval. Such a finding would suggest that maternal, endogenous resource allocation decisions are related to offspring physiology in a manner consistent with Kuzawas description. We observed exactly such a relationship among the Ache of Paraguay, suggesting that maternally mediated in utero signals of postnatal environments may be important determinants of later physiology. The implications of these findings are reviewed in light of life history and developmental plasticity theories and ourability to generalize the results to other populations. Recommendations for further empirical research are briefly summarized. Am. J. Hum. Biol., 2009.

Further new hominin fossils from the Kibish Formation, southwestern Ethiopia.

In addition to the new fragments of the Omo I skeleton, renewed fieldwork in the Kibish Formation along the lower reaches of the Omo River in southwestern Ethiopia has yielded new hominin finds from the Kibish Formation. The new finds include four heavily mineralized specimens: a partial left tibia and a fragment of a distal fibular diaphysis from Awokes Hominid Site (AHS), a parietal fragment, and a portion of a juvenile occipital bone. The AHS tibia and fibula derive from Member I and are contemporaneous with Omo I and II. The other specimens derive from Chads Hominid Site (CHS), and derive from either Member III or IV, which constrains their age between approximately 8.6 and approximately 104 ka.