Comparative ontogeny of functional aspects of human cervical vertebrae.

OBJECTIVES: Differences between adult humans and great apes in cervical vertebral morphology are well documented, but the ontogeny of this variation is still largely unexplored. This study examines patterns of growth in functionally relevant features of C1, C2, C4, and C6 in extant humans and apes to understand the development of their disparate morphologies. MATERIALS AND METHODS: Linear and angular measurements were taken from 530 cervical vertebrae representing 146 individual humans, chimpanzees, gorillas, and orangutans. Specimens were divided into three age-categories based on dental eruption: juvenile, adolescent, and adult. Inter- and intraspecific comparisons were evaluated using resampling methods. RESULTS: Of the eighteen variables examined here, seven distinguish humans from apes at the adult stage. Human-ape differences in features related to atlantoaxial joint function tend to be established by the juvenile stage, whereas differences in features related to the nuchal musculature and movement of the subaxial elements do not fully emerge until adolescence or later. The orientation of the odontoid process-often cited as a feature that distinguishes humans from apes-is similar in adult humans and adult chimpanzees, but the developmental patterns are distinct, with human adultlike morphology being achieved much earlier. DISCUSSION: The biomechanical consequences of the variation observed here is poorly understood. Whether the differences in growth patterns represent functional links to cranial development or postural changes, or both, requires additional investigation. Determining when humanlike ontogenetic patterns evolved in hominins may provide insight into the functional basis driving the morphological divergence between extant humans and apes.

New fossils of Australopithecus sediba reveal a nearly complete lower back.

Adaptations of the lower back to bipedalism are frequently discussed but infrequently demonstrated in early fossil hominins. Newly discovered lumbar vertebrae contribute to a near-complete lower back of Malapa Hominin 2 (MH2), offering additional insights into posture and locomotion in Australopithecus sediba. We show that MH2 possessed a lower back consistent with lumbar lordosis and other adaptations to bipedalism, including an increase in the width of intervertebral articular facets from the upper to lower lumbar column ('pyramidal configuration'). These results contrast with some recent work on lordosis in fossil hominins, where MH2 was argued to demonstrate no appreciable lordosis ('hypolordosis') similar to Neandertals. Our three-dimensional geometric morphometric (3D GM) analyses show that MH2's nearly complete middle lumbar vertebra is human-like in overall shape but its vertebral body is somewhat intermediate in shape between modern humans and great apes. Additionally, it bears long, cranially and ventrally oriented costal (transverse) processes, implying powerful trunk musculature. We interpret this combination of features to indicate that A. sediba used its lower back in both bipedal and arboreal positional behaviors, as previously suggested based on multiple lines of evidence from other parts of the skeleton and reconstructed paleobiology of A. sediba.

One of the defining features of humans is our ability to walk comfortably on two legs. To achieve this, our skeletons have evolved certain physical characteristics. For example, the lower part of the human spine has a forward curve that supports an upright posture; whereas the lower backs of chimpanzees and other apes ­ which walk around on four limbs and spend much of their time in trees ­ lack this curvature. Studying the fossilized back bones of ancient human remains can help us to understand how we evolved these features, and whether our ancestors moved in a similar way. Australopithecus sediba was a close-relative of modern humans that lived about two million years ago. In 2008, fossils from an adult female were discovered at a cave site in South Africa called Malapa. However, the fossils of the lower back region were incomplete, so it was unclear whether the female ­ referred to as Malapa Hominin 2 (MH2) ­ had a forward-curving spine and other adaptations needed to walk on two legs. Here, Williams et al. report the discovery of new A. sediba fossils from Malapa. The new fossils are mainly bones from the lower back, and they fit together with the previously discovered MH2 fossils, providing a nearly complete lower spine. Analysis of the fossils suggested that MH2 would have had an upright posture and comfortably walked on two legs, and the curvature of their lower back was similar to modern females. However, other aspects of the bones' shape suggest that as well as walking, A. sediba probably spent a significant amount of time climbing in trees. The findings of Williams et al. provide new insights in to our evolutionary history, and ultimately, our place in the natural world around us. Our lower back is prone to injury and pain associated with posture, pregnancy and exercise (or lack thereof). Therefore, understanding how the lower back evolved may help us to learn how to prevent injuries and maintain a healthy back.

Influences of passive intervertebral range of motion on cervical vertebral form.

OBJECTIVES: The cervical spine is the junction between the head and trunk, and it therefore facilitates head mobility and stability. The goal of this study is to test several predictions regarding cervical morphology and intervertebral ranges of motion. MATERIALS AND METHODS: Intervertebral ranges of motion for 12 primate species were collected via radiographs or taken from the literature. Morphometric data describing functionally relevant aspects of cervical vertebral morphology were obtained from museum specimens representing these species. We tested for correlations between intervertebral movement and vertebral form using phylogenetic generalized least-squares regression. RESULTS: Results demonstrate limited support for the hypothesis that range of motion (ROM) is influenced by cervical vertebral morphology. Few morphological variables correlate with ROM and no relationship is consistently significant across cervical joints. DISCUSSION: These results indicate that the relationship between vertebral morphology and joint ranges of motion is, at most, weak, providing little support the use of bony morphology to reconstruct axial mobility in fossil specimens. Future work should investigate the role of soft tissues in vertebral joint stability.

Comparative morphology and ontogeny of the thoracolumbar transition in great apes, humans, and fossil hominins.

Variation among extant hominoid taxa in the anatomy of the thoracolumbar vertebral transition is well-established and constitutes an important framework for making inferences about posture and locomotion in fossil hominins. However, little is known about the developmental bases of these differences, posing a challenge when interpreting the morphology of juvenile hominins. In this study, we investigated ontogenetic variation in the thoracolumbar transition of juvenile and adult great apes, humans, and fossils attributed to Australopithecus and early Pleistocene Homo erectus. For each vertebra involved in the transition, we quantified functionally relevant aspects of zygapophyseal form: facet curvature in the transverse plane, facet orientation relative to midline, and the shift in these variables across the thoracolumbar transition, from the antepenultimate rib-bearing thoracic to the first lumbar vertebra (L1). Among extant hominids, adult individuals of Pan and Homo exhibit a greater shift in facet morphology across the thoracolumbar transition in comparison to Gorilla and Pongo. This pattern is driven by interspecific differences in the L1 facets, with those of chimpanzees and humans being more curved and more sagittally oriented. Chimpanzees and humans also experience more change in facet morphology during development relative to gorillas and orangutans. Humans differ from chimpanzees in achieving their adultlike configuration much earlier in development. The fossil specimens indicate that early hominins had adult morphologies that were similar to those of extant Homo and Pan, and that they achieved their adult morphologies early in development, like extant humans. Although it is unclear why adult chimpanzees and hominins share an adult morphology, we speculate that the early acquisition of adultlike L1 zygapophyseal morphology in hominins is an evolutionary novelty related to conferring stability to a relatively long lumbar spine as young individuals are learning to walk bipedally.

Functional analyses of the primate upper cervical vertebral column.

Recent work has highlighted functional correlations between direct measures of head and neck posture and primate cervical bony morphology. Primates with more horizontal necks exhibit middle and lower cervical vertebral features that indicate increased mechanical advantage for deep nuchal musculature and mechanisms for column curvature formation and maintenance. How features of the C1 and C2 reflect quantified measures of posture have yet to be examined. This study incorporates bony morphology from the upper cervical levels from 20 extant primate species in order to investigate further how posture correlates with cervical vertebrae morphology. Results from phylogenetic generalized least-squares analyses indicate that few vertebral features exhibit a significant relationship with posture when accounting for differences in size. When size-adjusted traits were correlated with posture, vertebral variation had a stronger relationship with neck posture than head posture variables. Two C1 traits-relative posterior arch length and superior facet curvature-were correlated with neck posture variables. Relative posterior arch length exhibits a positive relationship with neck posture, while superior articular facet curvature demonstrates a negative relationship, such that as the neck becomes more horizontal, the greater the facet curvature. Four C2 features were also correlated with neck posture: relative pedicle and lamina lengths, relative superior facet orientation, and dens orientation. Relative pedicle and lamina lengths become craniocaudally longer as the neck becomes more horizontal. Relative C2 superior facet orientation and dens orientation exhibit negative correlations with posture, such that as the neck becomes more horizontal, the superior facet becomes more caudally inclined and the dens more dorsally inclined. These results produce a similar functional signal observed in the middle and lower cervical spine. Modeling the cervical vertebrae of more pronograde taxa within a sigmoidal spinal column model is further discussed and may prove useful in refining and testing future hypotheses of primate cervical mechanics.

Thoracic vertebral count and thoracolumbar transition in Australopithecus afarensis.

The evolution of the human pattern of axial segmentation has been the focus of considerable discussion in paleoanthropology. Although several complete lumbar vertebral columns are known for early hominins, to date, no complete cervical or thoracic series has been recovered. Several partial skeletons have revealed that the thoracolumbar transition in early hominins differed from that of most extant apes and humans. Australopithecus africanus, Australopithecus sediba, and Homo erectus all had zygapophyseal facets that shift from thoracic-like to lumbar-like at the penultimate rib-bearing level, rather than the ultimate rib-bearing level, as in most humans and extant African apes. What has not been clear is whether Australopithecus had 12 thoracic vertebrae as in most humans, or 13 as in most African apes, and where the position of the thoracolumbar transitional element was. The discovery, preparation, and synchrotron scanning of the Australopithecus afarensis partial skeleton DIK-1-1, from Dikika, Ethiopia, provides the only known complete hominin cervical and thoracic vertebral column before 60,000 years ago. DIK-1-1 is the only known Australopithecus skeleton to preserve all seven cervical vertebrae and provides evidence for 12 thoracic vertebrae with a transition in facet morphology at the 11th thoracic level. The location of this transition, one segment cranial to the ultimate rib-bearing vertebra, also occurs in all other early hominins and is higher than in most humans or extant apes. At 3.3 million years ago, the DIK-1-1 skeleton is the earliest example of this distinctive and unusual pattern of axial segmentation.

Functional morphology of the primate head and neck.

The vertebral column plays a key role in maintaining posture, locomotion, and transmitting loads between body components. Cervical vertebrae act as a bridge between the torso and head and play a crucial role in the maintenance of head position and the visual field. Despite its importance in positional behaviors, the functional morphology of the cervical region remains poorly understood, particularly in comparison to the thoracic and lumbar sections of the spinal column. This study tests whether morphological variation in the primate cervical vertebrae correlates with differences in postural behavior. Phylogenetic generalized least-squares analyses were performed on a taxonomically broad sample of 26 extant primate taxa to test the link between vertebral morphology and posture. Kinematic data on primate head and neck postures were used instead of behavioral categories in an effort to provide a more direct analysis of our functional hypothesis. Results provide evidence for a function-form link between cervical vertebral shape and postural behaviors. Specifically, taxa with more pronograde heads and necks and less kyphotic orbits exhibit cervical vertebrae with longer spinous processes, indicating increased mechanical advantage for deep nuchal musculature, and craniocaudally longer vertebral bodies and more coronally oriented zygapophyseal articular facets, suggesting an emphasis on curve formation and maintenance within the cervical lordosis, coupled with a greater resistance to translation and ventral displacement. These results not only document support for functional relationships in cervical vertebrae features across a wide range of primate taxa, but highlight the utility of quantitative behavioral data in functional investigations.

From the ground up: Integrative research in primate locomotion.

Primate locomotor adaptation and evolution is a principal and thriving area of research by biological anthropologists. Research in this field generally targets hypotheses regarding locomotor kinetics and kinematics, form-function associations in both the soft and hard tissue components of the musculoskeletal system, and reconstructing locomotor behavior in fossil primates. A wide array of methodological approaches is used to address adaptive hypotheses in all of these realms. Recent advances in three-dimensional shape capture, musculoskeletal physiological measurements, and analytical processing technologies (e.g., laser and CT-scans, 3D motion analysis systems, finite element analysis) have facilitated the collection and analysis of larger and more complex locomotor datasets than previously possible. With these advances in technology, new methods of form-function analyses can be developed to produce a more thorough understanding of how form reflects an organism's mechanical requirements, how shape is influenced by external environmental factors, and how these investigations of living taxa can inform questions of primate paleobiology. The papers in this special section of the American Journal of Physical Anthropology present research that builds on that foundation, by combining new data on living primates and new methodologies and approaches to answer a range of questions on extant and extinct primates.