A new turtle from the La Colonia Formation (Campanian–Maastrichtian), Patagonia,Argentina, with remarks on the evolution of the vertebral column in turtles

Abstract: Patagoniaemys gasparinae gen. et sp. nov. is a new stem turtle found in central Patagonia, Chubut Province, Argentina, in outcrops of the La Colonia Formation (Campanian–Maastrichtian). This is a turtle of relatively large size (carapace length c. 70 cm), and the preserved remains of the holotype consist of skull fragments and several postcranial elements including a nearly complete vertebral column. A phylogenetic analysis shows Patagoniaemys gasparinae gen. et sp. nov. forming a monophyletic group with Otwayemys cunicularius and Mongolochelys efremovi, as a sister group to Meiolaniidae. A comprehensive review confirms that formed cervical vertebrae appeared independently several times during turtle evolution: in the clade that includes Patagoniaemys gasparinae gen. et sp. nov. and Meiolaniidae, in some baenids, in the total group Pleurodira and in crown group Cryptodira. Likewise, formed caudal vertebrae appeared several times in turtle evolution.     

Functional aspects of strepsirrhine lumbar vertebral bodies and spinous processes

The relationship between form and function in the lumbar vertebral column has been well documented among platyrrhines and especially catarrhines, while functional studies of postcranial morphology among strepsirrhines have concentrated predominantly on the limbs. This morphometric study investigates biomechanically relevant attributes of the lumbar vertebral morphology of 20 species of extant strepsirrhines. With this extensive sample, our goal is to address the influence of positional behavior on lumbar vertebral form while also assessing the effects of body size and phylogenetic history. The results reveal distinctions in lumbar vertebral morphology among strepsirrhines in functional association with their habitual postures and primary locomotor behaviors. In general, strepsirrhines that emphasize pronograde posture and quadrupedal locomotion combined with leaping (from a pronograde position) have the relatively longest lumbar regions and lumbar vertebral bodies, features promoting sagittal spinal flexibility. Indrids and galagonids that rely primarily on vertical clinging and leaping with orthograde posture share a relatively short (i.e., stable and resistant to bending) lumbar region, although the length of individual lumbar vertebral bodies varies phylogenetically and possibly allometrically. The other two vertical clingers and leapers, Hapalemur and Lepilemur, more closely resemble the pronograde, quadrupedal taxa. The specialized, suspensory lorids have relatively short lumbar regions as well, but the lengths of their lumbar regions are influenced by body size, and Arctocebus has dramatically longer vertebral bodies than do the other lorids. Lumbar morphology among galagonids appears to reflect a strong phylogenetic signal superimposed on a functional one. In general, relative length of the spinous processes follows a positively allometric trend, although lorids (especially the larger-bodied forms) have relatively short spinous processes for their body size, in accordance with their positional repertoire. The results of the study broaden our understanding of postcranial adaptation in primates, while providing an extensive comparative database for interpreting vertebral morphology in fossil primates.     

Correlation between Hox code and vertebral morphology in archosaurs

The relationship between developmental genes and phenotypic variation is of central interest in evolutionary biology. An excellent example is the role of Hox genes in the anteroposterior regionalization of the vertebral column in vertebrates. Archosaurs (crocodiles, dinosaurs including birds) are highly variable both in vertebral morphology and number. Nevertheless, functionally equivalent Hox genes are active in the axial skeleton during embryonic development, indicating that the morphological variation across taxa is likely owing to modifications in the pattern of Hox gene expression. By using geometric morphometrics, we demonstrate a correlation between vertebral Hox code and quantifiable vertebral morphology in modern archosaurs, in which the boundaries between morphological subgroups of vertebrae can be linked to anterior Hox gene expression boundaries. Our findings reveal homologous units of cervical vertebrae in modern archosaurs, each with their specific Hox gene pattern, enabling us to trace these homologies in the extinct sauropodomorph dinosaurs, a group with highly variable vertebral counts. Based on the quantifiable vertebral morphology, this allows us to infer the underlying genetic mechanisms in vertebral evolution in fossils, which represents not only an important case study, but will lead to a better understanding of the origin of morphological disparity in recent archosaur vertebral columns.     

Adaptive landscapes challenge the “lateral-to-sagittal” paradigm for mammalian vertebral evolution

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Lumbar anomalies in the Shanidar 3 Neandertal

Recent examination of the Shanidar 3 remains revealed the presence of anomalous bilateral arthroses in the lumbar region. This paper describes this developmental anomaly, as well as several degenerative changes and offers potential etiologies.The Shanidar 3 remains represent an adult male Neandertal, approximately 35–50 years of age, dating to the Last Glacial. Although the partial skeleton is fragmentary, preserved elements include an almost complete set of ribs, portions of all thoracic vertebrae, all lumbar vertebrae, and the sacrum. Vertebral articulations from S1–T1 can be confidently assigned. The vertebra designated L1 is well preserved but lacks transverse processes. Instead, well defined bilateral articular surfaces, rather than transverse processes, are located on the pedicles. The skeletal elements associated with the anomalous L1 articulations were not recovered.The most likely interpretation is that the arthroses in question represent the facets for a 13th pair of ribs, a rare condition in modern hominid populations. Such lumbar developmental anomalies are an infrequent expression of a larger complex of cranial-caudal border shifting seen in the vertebral column. These shifts result in a change in the usual boundaries between the distinctive vertebral regions and are responsible for the majority of variability present in the vertebral column.     

Sex differences in the vertebral column of gombe chimpanzees

Male and female chimpanzees from Gombe National Park, Tanzania (Pan troglodytes schweinfurthii) differ in live body weights but not in cranial capacity or fore-and hindlimb long bone lengths. Skeletal dimensions of the limbs and vertebral column indicate a mosaic of sex differences. Vertebral column measurements generally are greater in males. While linear measurements identify differences in the breadth and depth of the lower cervical and upper thoracic vertebrae, areal assessments show significant differences in weight-bearing surfaces throughout the thoracic and lumbar segments. These can be interpreted in terms of distribution of weight and body composition (i.e. amount of musculature).     

Pathobiomechanical impairments of the vertebral column in intervertebral disk protrusion and herniation

Magnetic resonance imaging of patients with intervertebral disk herniation was used to study the stages of degenerative and dystrophic processes in the spinal structures in the presence of intervertebral disk protrusion and herniation. Differences were found in the pathobiomechanical mechanisms in the spinal motor segments of herniation and protrusion in the area of their localization and in the adjacent spinal motor segments. Among the symptoms traditionally analyzed, joint facet joint arthritis and arthrosoarthritis classified as spondyloarthritis by radiodiagnosis were examined for their impact on the rate of herniation and protrusion.     

Morphological correlates of tail length in the catarrhine sacrum

Taillessness is a distinctive synapomorphy of the Hominoidea that has implications for interpretation of the locomotor behaviors and phylogenetic affinities of the clade’s earliest members. However, difficulties persist in confidently identifying taillessness in the catarrhine fossil record, stemming largely from our limited knowledge of the anatomical features with which the tail is associated. Here, we compare the morphology of the sacrum, the sole bony link between the tail and the rest of the body, among extant tailless hominoids and a broad sample of extant cercopithecoids known to vary in tail length (i.e., ‘very short’, ‘short’, and ‘long’) in order to identify morphological correlates of tail length. We examine three features of the sacrum, including the shape of the sacrum’s caudal articular surface (CAS), the sacrocaudal articulation (SCA) angle, and the lateral expansion of the last sacral vertebra’s transverse processes. Compared with all other taxa, ‘long’-tailed cercopithecoids have significantly more circularly-shaped CASs, more acute SCA angles, and more laterally expanded transverse processes of the last sacral vertebra. Tailless hominoids have significantly more elliptically-shaped CASs and less laterally expanded transverse processes than all tailed cercopithecoids, but in the latter parameter, they only differ significantly from ‘long’-tailed cercopithecoids. Cercopithecoids with ‘short’ and ‘very short’ tails are intermediate between tailless hominoids and ‘long’-tailed cercopithecoids with respect to CAS shape and lateral expansion of the transverse processes. SCA angle distinguishes clearly among all three cercopithecoid tail length groups. The results of this study provide evidence for significant differences in sacral morphology among extant catarrhines known to differ in tail length, and have implications for making inferences about tail length and function in extinct catarrhines.     

Spiral computed tomography in the diagnosis of inflammatory diseases of the vertebral column

A Somatom Plus 4 spiral computed tomograph was used to examine 40 patients with purulent spondylitis and 8 patients with tuberculous spondylitis. Analysis of the primary examination, postoperative status, and follow-up of patients on medical therapy identified the main signs of an inflammatory process from a great variety of the computed tomographic (CT) signs of a lesion. They made it possible not only to reveal the inflammatory nature of a lesion, but also to differentiate purulent and tuberculous spondylitis. The main CT signs that allow a purulent spondylitis to be differentiated from tuberculous one are as follows: the location of involved veterbrae, the number of and the pattern of destruction of trabecular and cortical layers, the extent of and the form of paravertebral tissue lesion.     

Variability in mammalian dentitions: size-related bias in the coefficient of variation

The coefficient of variation has been used in many evolutionary studies. However, a strong negative correlation between this index and size may artificially inflate the apparent variability of small traits. This is most pronounced when variables whose size differs by more than an order of magnitude are compared or when the index is applied to variables whose size is within an order of magnitude of their measurement error. When this is likely to affect conclusions, other indexes of variability should be considered. One alternative is to use the standard deviation of log-transformed data; however, this index is sometimes still correlated with variable size so care should be exercised in its use. Another alternative is to regress the standard deviation onto mean variable size; however, this method may also be misleading if variables are not randomly distributed about the regression line. As an example of the effect of bias in the coefficient of variation, previous studies of mammalian dental variability profiles were re-evaluated. It was found that variation among teeth is relatively homogenous, both within and among species. The exception is that the canines of some mammalian species have variability that is considerably higher than would be expected from tooth size. Previous explanations of variability patterns that invoked developmental fields are incompatible with the new data.     

Strong links between genomic and anatomical diversity in both mammalian olfactory chemosensory systems

Mammalian olfaction comprises two chemosensory systems: the odorant-detecting main olfactory system (MOS) and the pheromone-detecting vomeronasal system (VNS). Mammals are diverse in their anatomical and genomic emphases on olfactory chemosensation, including the loss or reduction of these systems in some orders. Despite qualitative evidence linking the genomic evolution of the olfactory systems to specific functions and phenotypes, little work has quantitatively tested whether the genomic aspects of the mammalian olfactory chemosensory systems are correlated to anatomical diversity. We show that the genomic and anatomical variation in these systems is tightly linked in both the VNS and the MOS, though the signature of selection is different in each system. Specifically, the MOS appears to vary based on absolute organ and gene family size while the VNS appears to vary according to the relative proportion of functional genes and relative anatomical size and complexity. Furthermore, there is little evidence that these two systems are evolving in a linked fashion. The relationships between genomic and anatomical diversity strongly support a role for natural selection in shaping both the anatomical and genomic evolution of the olfactory chemosensory systems in mammals.     

Domains within the mammalian ornithine decarboxylase messenger RNA have evolved independently and episodically

Summary Ornithine decarboxylase (ODC) is the first enzyme in the polyamine biosynthetic pathway. We have studied the evolutionary history of the mammalian ODC mRNA, focusing on the rate of accumulation of sequence divergence within specific subregions of the molecule. The phylogenetic relationships among the mRNAs from several mammalian species, including two mouse species, rat, hamster, and human, were determined based upon the numbers of synonymous substitutions in pair-wise comparisons of mRNA coding regions. The separation times for the mRNAs were very similar to those for the corresponding species, suggesting that ODC is encoded by orthologous genes in the different species. Analysis of divergence patterns in four subregions, or domains, of the mRNA (the 5untranslated region, the coding region, and two domains of the 3-untranslated region) showed that the domains have evolved in a noncoordinate fashion. Furthermore, evolution of each subregion has been episodic, with periods of both rapid and slow sequence divergence. We suggest that the episodic pattern of ODC mRNA evolution may indicate the existence of selection pressures that were exerted in a time- and domain-specific manner during mammalian speciation.     

Dexamethasone stimulates osteogenic differentiation in vertebral and femoral bone marrow cell cultures: Comparison of IGF-I gene expression

Osteoblast-like cell cultures have been established from the marrow of adult rat vertebrae. We have simultaneously examined the response to dexamethasone (dex) treatment in cultures of young adult female vertebral and femoral marrow cells. Alkaline phosphatase (AP) activity was analyzed as well as the expression of mRNAs for osteocalcin (OC) and insulin-like growth factor I (IGF-I). The vertebral and femoral marrow cells were maintained for 7 days in primary culture with or without 10⁻⁸ M dex and then 6 days in secondary culture without dex or with 10⁻⁸ M or 10⁻⁷ M dex. All cells were examined on day 6 of secondary culture. Vertebral and femoral cultures each expressed the highest AP enzyme levels when grown with dex in primary culture (10⁻⁸ M) and secondary culture (10⁻⁷ M). Under all experimental conditions, vertebral cultures had lower AP enzyme activity than femoral cultures. When dex was omitted from secondary culture, OC gene expression was not detected in either vertebral or femoral passaged cells even if dex was present in primary culture. For dex conditions where OC was expressed, vertebral cultures had higher OC mRNA steady-state levels than femoral cultures. IGF-I gene expression was detected by Northern analysis in both vertebral and femoral secondary cultures. However, vertebral marrow cultures had much higher IGF-I mRNA levels compared to femoral cultures whether or not dex was present in primary culture. These findings demonstrate that dex supports the differentiation of both vertebral and femoral adult marrow osteogenic cells into osteoblasts. Our results support the hypothesis that osteoblastic marrow cultures differ depending upon which location in the skeleton they are from and that there are skeletal site–dependent differences in the insulin-like growth factor system components. J. Cell. Biochem. 71:382–391, 1998. © 1998 Wiley-Liss, Inc.     

Radiological evaluation of bone status in the jaw and in the vertebral column in a group of women

Vertebral bone mineral content was determined in a group of 56 women, ages 30–62. These measurements were compared with the status of supporting bone in the jaws (alveolar, molar and bicuspid) and with gingival health. There was a significant decline in vertebral bone mineral content from the pre- to post-menopausal group. Molar and bicuspid measurements were highly correlated. There was some association between lumbar bone mineral content and molar bone status for postmenopausal women. For postmenopausal women, the cases of greatest percent bone loss in alveolar crest were associated with lower lumbar bone mineral content. Gingival health did not confound the bone status measurements. The 56 subjects did not exhibit the degree of reduction in bone density that is observed in the general population. Further investigation using these radiographic techniques may reveal a link between substantial bone loss in the jaw and moderate to severe bone loss in the lumbar vertebrae.     

Brain evolution and development: adaptation,allometry and constraint

Phenotypic traits are products of two processes: evolution and development. But how do these processes combine to produce integrated phenotypes? Comparative studies identify consistent patterns of covariation, or allometries, between brain and body size, and between brain components, indicating the presence of significant constraints limiting independent evolution of separate parts. These constraints are poorly understood, but in principle could be either developmental or functional. The developmental constraints hypothesis suggests that individual components (brain and body size, or individual brain components) tend to evolve together because natural selection operates on relatively simple developmental mechanisms that affect the growth of all parts in a concerted manner. The functional constraints hypothesis suggests that correlated change reflects the action of selection on distributed functional systems connecting the different sub-components, predicting more complex patterns of mosaic change at the level of the functional systems and more complex genetic and developmental mechanisms. These hypotheses are not mutually exclusive but make different predictions. We review recent genetic and neurodevelopmental evidence, concluding that functional rather than developmental constraints are the main cause of the observed patterns.     

Structure and function in mammalian societies

Traditional interpretations of the evolution of animal societies have suggested that their structure is a consequence of attempts by individuals to maximize their inclusive fitness within constraints imposed by their social and physical environments. In contrast, some recent re-interpretations have argued that many aspects of social organization should be interpreted as group-level adaptations maintained by selection operating between groups or populations. Here, I review our current understanding of the evolution of mammalian societies, focusing, in particular, on the evolution of reproductive strategies in societies where one dominant female monopolizes reproduction in each group and her offspring are reared by other group members. Recent studies of the life histories of females in these species show that dispersing females often have little chance of establishing new breeding groups and so are likely to maximize their inclusive fitness by helping related dominants to rear their offspring. As in eusocial insects, increasing group size can lead to a progressive divergence in the selection pressures operating on breeders and helpers and to increasing specialization in their behaviour and life histories. As yet, there is little need to invoke group-level adaptations in order to account for the behaviour of individuals or the structure of mammalian groups.     

Uncertainty analysis of material properties and morphology parameters in numerical models regarding the motion of lumbar vertebral segments

Abstract

The kinematics of a spinal motion segment is determined by the material properties of the soft-tissue and the morphology. The material properties can vary within subjects and between vertebral levels, leading to a wide possible range of motion of a spinal segment independently on its morphology. The goal of this numerical study was to identify the most influential material parameters concerning the kinematics of a spinal motion segment and their plausible ranges. Then, a method was tested to deduce the material properties automatically, based on a given ROM and morphology. A fully parametric finite element model of the morphology and material properties of a lumbar spinal motion segment was developed. The impact of uncertainty of twelve spinal material parameters, as well as the size of the gap between the articular surfaces of the facet joints was examined. The simulation results were compared to our own in vitro data. The flexibility of a lumbar segment was especially influenced by the properties of the anterior annulus region, the facet gap size and the interspinous ligament. The high degree of uncertainty in the material properties and facet gap size published in the literature can lead to a wide scatter in the motion of a spinal segment, with a range of 6°-17° in the intact condition in flexion/extension, from 5°-22° in lateral bending and from 3°-14° in axial rotation. Statistical analysis of the variability might help to estimate the sensitivity and total uncertainty propagated through biomechanical simulations, affecting the reliability of the predictions.