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.

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.

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.

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.