The cervical spine of the American barn owl (Tyto furcata pratincola): I. Anatomy of the vertebrae and regionalization in their S-shaped arrangement.
ABSTRACT: BACKGROUND: Owls possess an extraordinary neck and head mobility. To understand this mobility it is necessary to have an anatomical description of cervical vertebrae with an emphasis on those criteria that are relevant for head positioning. No functional description specific to owls is available. METHODOLOGY/PRINCIPAL FINDINGS: X-ray films and micro-CT scans were recorded from American barn owls (Tyto furcata pratincola) and used to obtain three-dimensional head movements and three-dimensional models of the 14 cervical vertebrae (C1-C14). The diameter of the vertebral canal, the zygapophyseal protrusion, the distance between joint centers, and the pitching angle were quantified. Whereas the first two variables are purely osteological characteristics of single vertebrae, the latter two take into account interactions between vertebrae. These variables change in characteristic ways from cranial to caudal. The vertebral canal is wide in the cranial and caudal neck regions, but narrow in the middle, where both the zygapophyseal protrusion and the distance between joint centers are large. Pitching angles are more negative in the cranial and caudal neck regions than in the middle region. Cluster analysis suggested a complex regionalization. Whereas the borders (C1 and C13/C14) formed stable clusters, the other cervical vertebrae were sorted into 4 or 5 additional clusters. The borders of the clusters were influenced by the variables analyzed. CONCLUSIONS/SIGNIFICANCE: A statistical analysis was used to evaluate the regionalization of the cervical spine in the barn owl. While earlier measurements have shown that there appear to be three regions of flexibility of the neck, our indicators suggest 3-7 regions. These many regions allow a high degree of flexibility, potentially facilitating the large head turns that barn owls are able to make. The cervical vertebral series of other species should also be investigated using statistical criteria to further characterize morphology and the potential movements associated with it.
Project description:Several evolutionary theories have been proposed to explain the adaptation of the long giraffe neck; however, few studies examine the fossil cervical vertebrae. We incorporate extinct giraffids, and the okapi and giraffe cervical vertebral specimens in a comprehensive analysis of the anatomy and elongation of the neck. We establish and evaluate 20 character states that relate to general, cranial and caudal vertebral lengthening, and calculate a length-to-width ratio to measure the relative slenderness of the vertebrae. Our sample includes cervical vertebrae (n=71) of 11 taxa representing all seven subfamilies. We also perform a computational comparison of the C3 of Samotherium and Giraffa camelopardalis, which demonstrates that cervical elongation occurs disproportionately along the cranial-caudal vertebral axis. Using the morphological characters and calculated ratios, we propose stages in cervical lengthening, which are supported by the mathematical transformations using fossil and extant specimens. We find that cervical elongation is anisometric and unexpectedly precedes Giraffidae. Within the family, cranial vertebral elongation is the first lengthening stage observed followed by caudal vertebral elongation, which accounts for the extremely long neck of the giraffe.
Project description:Mammals show a very low level of variation in vertebral count, particularly in the neck. Phenotypes exhibited at various stages during the development of the axial skeleton may play a key role in testing mechanisms recently proposed to explain this conservatism. Here, we provide osteogenetic data that identify developmental criteria with which to recognize cervical vs. noncervical vertebrae in mammals. Except for sloths, all mammals show the late ossification of the caudal-most centra in the neck after other centra and neural arches. In sloths with 8-10 ribless neck vertebrae, the caudal-most neck centra ossify early, matching the pattern observed in cranial thoracic vertebrae of other mammals. Accordingly, we interpret the ribless neck vertebrae of three-toed sloths caudal to V7 as thoracic based on our developmental criterion. Applied to the unusual vertebral phenotype of long-necked sloths, these data support the interpretation that elements of the axial skeleton with origins from distinct mesodermal tissues have repatterned over the course of evolution.
Project description:Giraffidae are represented by many extinct species. The only two extant taxa possess diametrically contrasting cervical morphology, as the okapi is short-necked and the giraffe is exceptionally long-necked. Samotherium major, known from the Late Miocene of Samos in Greece and other Eurasian localities, is a key extinct giraffid; it possesses cervical vertebrae that are intermediate in the evolutionary elongation of the neck. We describe detailed anatomical features of the cervicals of S. major, and compare these characteristics with the vertebrae of the two extant giraffid taxa. Based on qualitative morphological characters and a quantitative analysis of cervical dimensions, we find that the S. major neck is intermediate between that of the okapi and the giraffe. Specifically, the more cranial (C2-C3) vertebrae of S. major represent a mosaic of features shared either with the giraffe or with the okapi. The more caudal (C5-C7) S. major vertebrae, however, appear transitional between the two extant taxa, and hence are more unique. Notably, the C6 of S. major exhibits a partially excavated ventral lamina that is strong cranially but completely absent on the caudal half of the ventral vertebral body, features between those seen in the giraffe and the okapi. Comprehensive anatomical descriptions and measurements of the almost-complete cervical column reveal that S. major is a truly intermediate-necked giraffid. Reconstructions of the neck display our findings.
Project description:<h4>Background</h4>Birds have highly mobile necks, but neither the details of how they realize complex poses nor the evolution of this complex musculoskeletal system is well-understood. Most previous work on avian neck function has focused on dorsoventral flexion, with few studies quantifying lateroflexion or axial rotation. Such data are critical for understanding joint function, as musculoskeletal movements incorporate motion around multiple degrees of freedom simultaneously. Here we use biplanar X-rays on wild turkeys to quantify three-dimensional cervical joint range of motion in an avian neck to determine patterns of mobility along the cranial-caudal axis.<h4>Results</h4>Range of motion can be generalized to a three-region model: cranial joints are ventroflexed with high axial and lateral mobility, caudal joints are dorsiflexed with little axial rotation but high lateroflexion, and middle joints show varying amounts axial rotation and a low degree of lateroflexion. Nonetheless, variation within and between regions is high. To attain complex poses, substantial axial rotation can occur at joints caudal to the atlas/axis complex and zygapophyseal joints can reduce their overlap almost to osteological disarticulation. Degrees of freedom interact at cervical joints; maximum lateroflexion occurs at different dorsoventral flexion angles at different joints, and axial rotation and lateroflexion are strongly coupled. Further, patterns of joint mobility are strongly predicted by cervical morphology.<h4>Conclusion</h4>Birds attain complex neck poses through a combination of mobile intervertebral joints, coupled rotations, and highly flexible zygapophyseal joints. Cranial-caudal patterns of joint mobility are tightly linked to cervical morphology, such that function can be predicted by form. The technique employed here provides a repeatable protocol for studying neck function in a broad array of taxa that will be directly comparable. It also serves as a foundation for future work on the evolution of neck mobility along the line from non-avian theropod dinosaurs to birds.
Project description:Sauropods are often imagined to have held their heads high atop necks that ascended in a sweeping curve that was formed either intrinsically because of the shape of their vertebrae, or behaviorally by lifting the head, or both. Their necks are also popularly depicted in life with poses suggesting avian flexibility. The grounds for such interpretations are examined in terms of vertebral osteology, inferences about missing soft tissues, intervertebral flexibility, and behavior. Osteologically, the pronounced opisthocoely and conformal central and zygapophyseal articular surfaces strongly constrain the reconstruction of the cervical vertebral column. The sauropod cervico-dorsal vertebral column is essentially straight, in contrast to the curvature exhibited in those extant vertebrates that naturally hold their heads above rising necks. Regarding flexibility, extant vertebrates with homologous articular geometries preserve a degree of zygapophyseal overlap at the limits of deflection, a constraint that is further restricted by soft tissues. Sauropod necks, if similarly constrained, were capable of sweeping out large feeding surfaces, yet much less capable of retracting the head to explore the enclosed volume in an avian manner. Behaviorally, modern vertebrates generally assume characteristic neck postures which are close to the intrinsic curvature of the undeflected neck. With the exception of some vertebrates that can retract their heads to balance above their shoulders at rest (e.g., felids, lagomorphs, and some ratites), the undeflected neck generally predicts the default head height at rest and during locomotion.
Project description:The objectives of this observational, cross-sectional study were to characterize and establish the prevalence of osseous proliferation of articular surfaces, joint margins and adjacent soft tissue attachments (i.e., joint capsule and deep spinal muscles) in a mixed population of horses of variable ages, sizes, and breeds to better capture the full spectrum of disease affecting the cervical articular processes. Cranial and caudal articular processes of the cervical and first three thoracic vertebrae (C2-T3) from 55 horses without a primary complaint of neck pain were evaluated for the presence and severity of abnormal bony changes. Data were analyzed to compare alterations in joint margin quadrants, paired articular surfaces within a synovial articulation, left-right laterality, and vertebral level distributions and to determine associations with age, wither height and sex. Seventy-two percent of articular processes had bony changes that were considered abnormal. Osteophyte formation was the most common bony change noted. Overall grades of severity included: normal (28%), mild (45%), moderate (22%), and severe (5%). The highest prevalence of mild changes was localized to the C3-C6 vertebral levels; moderate changes to C6-T2; and severe changes to C2-C3 and C6-T2. Most paired articular surfaces and left-right grades of severity were not significantly different. The grade of osseous pathology was positively associated with both age and wither height. A high prevalence and wide variety of abnormal bony changes of varying severity were found in articular processes across all vertebral levels. The clinical significance of the described lesions is unknown, but the findings are expected to enhance the reporting of articular process and periarticular changes noted on advanced diagnostic imaging of the equine cervical and cranial thoracic vertebral regions.
Project description:PMMA is the most common bone substitute used for vertebroplasty. An increased fracture rate of the adjacent vertebrae has been observed after vertebroplasty. Decreased failure strength has been noted in a laboratory study of augmented functional spine units (FSUs), where the adjacent, non-augmented vertebral body always failed. This may provide evidence that rigid cement augmentation may facilitate the subsequent collapse of the adjacent vertebrae. The purpose of this study was to evaluate whether the decrease in failure strength of augmented FSUs can be avoided using low-modulus PMMA bone cement. In cadaveric FSUs, overall stiffness, failure strength and stiffness of the two vertebral bodies were determined under compression for both the treated and untreated specimens. Augmentation was performed on the caudal vertebrae with either regular or low-modulus PMMA. Endplate and wedge-shaped fractures occurred in the cranial and caudal vertebrae in the ratios endplate:wedge (cranial:caudal): 3:8 (5:6), 4:7 (7:4) and 10:1 (10:1) for control, low-modulus and regular cement group, respectively. The mean failure strength was 3.3 +/- 1 MPa with low-modulus cement, 2.9 +/- 1.2 MPa with regular cement and 3.6 +/- 1.3 MPa for the control group. Differences between the groups were not significant (p = 0.754 and p = 0.375, respectively, for low-modulus cement vs. control and regular cement vs. control). Overall FSU stiffness was not significantly affected by augmentation. Significant differences were observed for the stiffness differences of the cranial to the caudal vertebral body for the regular PMMA group to the other groups (p < 0.003). The individual vertebral stiffness values clearly showed the stiffening effect of the regular cement and the lesser alteration of the stiffness of the augmented vertebrae using the low-modulus PMMA compared to the control group (p = 0.999). In vitro biomechanical study and biomechanical evaluation of the hypothesis state that the failure strength of augmented functional spine units could be better preserved using low-modulus PMMA in comparison to regular PMMA cement.
Project description:Sloths are one of only two exceptions to the mammalian 'rule of seven' vertebrae in the neck. As a striking case of breaking the evolutionary constraint, the explanation for the exceptional number of cervical vertebrae in sloths is still under debate. Two diverging hypotheses, both ultimately linked to the low metabolic rate of sloths, have been proposed: hypothesis 1 involves morphological transformation of vertebrae due to changes in the Hox gene expression pattern and hypothesis 2 assumes that the Hox gene expression pattern is not altered and the identity of the vertebrae is not changed. Direct evidence supporting either hypothesis would involve knowledge of the vertebral Hox code in sloths, but the realization of such studies is extremely limited. Here, on the basis of the previously established correlation between anterior Hox gene expression and the quantifiable vertebral shape, we present the morphological regionalization of the neck in three different species of sloths with aberrant cervical count providing indirect insight into the vertebral Hox code.Shape differences within the cervical vertebral column suggest a mouse-like Hox code in the neck of sloths. We infer an anterior shift of HoxC-6 expression in association with the first thoracic vertebra in short-necked sloths with decreased cervical count, and a posterior shift of HoxC-5 and HoxC-6 expression in long-necked sloths with increased cervical count.Although only future developmental analyses in non-model organisms, such as sloths, will yield direct evidence for the evolutionary mechanism responsible for the aberrant number of cervical vertebrae, our observations lend support to hypothesis 1 indicating that the number of modules is retained but their boundaries are displaced. Our approach based on quantified morphological differences also provides a reliable basis for further research including fossil taxa such as extinct 'ground sloths' in order to trace the pattern and the underlying genetic mechanisms in the evolution of the vertebral column in mammals.
Project description:Aegicetus gehennae is a new African protocetid whale based on a partial skull with much of an associated postcranial skeleton. The type specimen, Egyptian Geological Museum, Cairo [CGM] 60584, was found near the base of the early-Priabonian-age (earliest late Eocene) Gehannam Formation of the Wadi Al Hitan World Heritage Site in Egypt. The cranium is distinctive in having ventrally-deflected exoccipitals. The vertebral column is complete from cervical C1 through caudal Ca9, with a vertebral formula of 7:15:4:4:9+, representing, respectively, the number of cervical, thoracic, lumbar, sacral, and caudal vertebrae. CGM 60584 has two more rib-bearing thoracic vertebrae than other known protocetids, and two fewer lumbars. Sacral centra are unfused, and there is no defined auricular surface on the ilium. Thus there was no weight-bearing sacroiliac joint. The sternum is distinctive in being exceptionally broad and flat. The body weight of CGM 60584, a putative male, is estimated to have been about 890 kg in life. Long bones of the fore and hind limbs are shorter than expected for a protocetid of this size. Bones of the manus are similar in length and more robust compared to those of the pes. A log vertebral length profile for CGM 60584 parallels that of middle Eocene Maiacetus inuus through the anterior and middle thorax, but more posterior vertebrae are proportionally longer. Vertebral elongation, loss of a sacroiliac articulation, and hind limb reduction indicate that Aegicetus gehennae was more fully aquatic and less specialized as a foot-powered swimmer than earlier protocetids. It is doubtful that A. gehennae had a tail fluke, and the caudal flattening known for basilosaurids is shorter relative to vertebral column length than flattening associated with a fluke in any modern whale. Late protocetids and basilosaurids had relatively long skeletons, longer than those known earlier and later, and the middle-to-late Eocene transition from foot-powered to tail-powered swimming seemingly involved some form of mid-body-and-tail undulation.
Project description:Vertebral number is the most variable trait among vertebrates. In addition to the vertebral number, the ratio of abdominal to caudal vertebrae is a variable trait. The vertebral number and the ratio of abdominal to caudal vertebrae contribute to vertebrate diversity. It is very interesting to know how to determine the vertebral number and the ratio of abdominal to caudal vertebrae. In this study, we identify differences in the vertebral number and the ratio of abdominal vertebrae to vertebral number between two inbred lines of medaka, namely, Hd-rRII1 and Kaga. To identify the genetic factor of those differences, we performed quantitative trait locus (QTL) analysis for vertebral number and the ratio of abdominal vertebrae to vertebral number using 200 F(2) fish. Our results show a suggestive QTL of the ratio of abdominal vertebrae to vertebral number on chromosome 15, and five QTL of vertebral number on chromosomes 1, 10, 11, 17, and 23. The QTL on chromosome 15 contains hoxDb cluster genes. The QTL of vertebral number include some genes related to the segmentation clock and axial elongation. In addition, we show that the difference in vertebral number between two inbred lines is derived from differences in the anteroposterior length of somites. Our results emphasize that the developmental process should be considered in genetic analyses for vertebral number.