Effects of annular cranial vault modification on the cranial base and face

Artificial modification of the cranial vault was practiced by a number of prehistoric and protohistoric populations, frequently during an infant's first year of life. We test the hypothesis that, in addition to its direct effects on the cranial vault, annular cranial vault modification has a significant indirect effect on cranial base and facial morphology. Two skeletal series from the Pacific Northwest Coast, which include both nonmodified and modified crania, were used: the Kwakiutl (62 nonmodified, 45 modified) and Nootka (28 nonmodified, 20 modified). Three-dimensional coordinates of 53 landmarks were obtained using a diagraph, and 36 landmarks were used to define nine finite elements in the cranial vault, cranial base, and face. Finite element scaling was used to compare average nonmodified and average modified crania, and the significance of the results were evaluated using a bootstrap test. Annular modification of the cranial vault produces significant effects on the morphology of the cranial base and face. Annular modification in the Kwakiutl resulted in restrictions of the cranial vault in the medial-lateral and superior-inferior dimensions and an increase in anterior-posterior growth. Similar dimensional changes are observed in the cranial base. The Kwakiutl face is increased anterior-posteriorly and reduced anterior-laterally to posterior-medially. Similar effects of modification are observed in the Nootka cranial vault and cranial base, though not in the face. These results demonstrate the developmental interdependence of the cranial vault, cranial base, and face. © 1993 Wiley-Liss, Inc.     

Effects of fronto-occipital artificial cranial vault modification on the cranial base and face.

Artificial reshaping of the cranial vault has been practiced by many human groups and provides a natural experiment in which the relationships of neurocranial, cranial base, and facial growth can be investigated. We test the hypothesis that fronto-occipital artificial reshaping of the neurocranial vault results in specific changes in the cranial base and face. Fronto-occipital reshaping results from the application of pads or a cradle board which constrains cranial vault growth, limiting growth between the frontal and occipital and allowing compensatory growth of the parietals in a mediolateral direction. Two skeletal series including both normal and artificially modified crania are analyzed, a prehistoric Peruvian Ancon sample (47 normal, 64 modified crania) and a Songish Indian sample from British Columbia (6 normal, 4 modified). Three-dimensional coordinates of 53 landmarks were measured with a diagraph and used to form 9 finite elements as a prelude to finite element scaling analysis. Finite element scaling was used to compare average normal and modified crania and the results were evaluated for statistical significance using a bootstrap test. Fronto-occipitally reshaped Ancon crania are significantly different from normal in the vault, cranial base, and face. The vault is compressed along an anterior-superior to posterior-inferior axis and expanded along a mediolateral axis in modified individuals. The cranial base is wider and shallower in the modified crania and the face is foreshortened and wider with the anterior orbital rim moving inferior and posterior towards the cranial base. The Songish crania display a different modification of the vault and face, indicating that important differences may exist in the morphological effects of fronto-occipital reshaping from one group to another.     

Maxillary changes and occlusal traits in crania with artificial fronto-occipital deformation

Artificial fronto-occipital deformation of the cranial vault was typical of pre-Columbian cultures in the central Andean coastal regions. We have studied the influence of this deformation on maxillary and mandibular morphology. Measurements were performed on 86 adult Ancon skulls with anteroposterior deformation. Undeformed skulls from the area of Makatampu (n = 52) were used as the control group. To explore the influence of the deformity on occlusion, the skulls were categorized using the Angle classification and the alignment of the interincisor midline. In the group of deformed skulls, there was an increase in lateral growth of the vault and of the base of the skull (P < 0.001), giving rise to a greater interpterygoid width of the maxilla (P < 0.001), and an increase in the transverse diameter of the palatal vault. The mandible presented an increase in the length of the rami (P < 0.001) and in the intercondylar width, with no alteration of mandibular length. The deformed skulls had normal (class I) occlusion, with no displacement of the midline. The difference in the asymmetry index between the two groups was not statistically significant. Artificial fronto-occipital deformation of the cranial vault provoked compensatory lateral expansion of the base that was correlated with the transverse development of the maxilla and mandible. Occlusion and sagittal intermaxillary position were not affected by the cranial deformity. These results provide evidence of the integration between the neurocranium and the viscerocranium in craniofacial development, and support the hypothesis of a compensatory effect of function.     

Cranial capacity/cranial base relationships and prediction of vault form: A canonical correlation analysis

Canonical correlation analysis was used to test an hypothesized morphological relationship between vault form and cranial capacity relative to length of the chondrocranium. Ninety-five adult male Czech skulls were measured for vault form expressed as length, width and height of the brain case; the chondrocranium was represented by nasion-basion and basion-opisthion lengths. In terms of explained variation, the first and most important dimension of covariation between vault and chondrocranial variables was size. The second most significant dimension of covariation expressed the hypothesized shape relationships—i.e., overall size being equal, the shorter the chondrocranial base relative to cranial capacity, the shorter and wider the vault. Furthermore, the competing hypothesis that vault form is determined by facial length proved untenable since facial length was predictive of vault shape only when measured as prosthion-basion, a measure that incorporates basal length. When corrected for basal length, facial length is unrelated to vault form. The results are consistent with the assumption that phylogenetic and microevolutionary trends toward brachycephaly in man stem from changes in the relationship between two components of skull growth, the chondrocranial base and the brain.     

Craniofacial changes in hemifacial microsomia

Cephalometry, X-ray cephalometry, and somatoscopy were used in the studies of 65 adult males with a severe unilateral microtia subdivided into three groups: with marked asymmetry, with slight asymmetry, and without any obvious facial asymmetry. The group with marked asymmetries was designated as hemifacial microsomia. In this group the affected side of the face was depressed on the average from above and from below towards the level of the external auditory canal. The center of the anteroposterior reduction was situated in the region in front of the pterygomaxillar fissure. The anteroposterior and vertical facial dimensions on the affected side were reduced most markedly, while the width dimensions showed the slightest changes. Hypoplasia was most severe within the lower face and increased towards the otocephalic centre. The mandibular joint was displaced in an anteroinferior and medial direction. Hypertelorism did not occur, but the orbit on the affected side was smaller in height and was frequently vertically dislocated. The facial profile was unchanged except for retrusion of the chin and increased frequency of bite disorders. The mobility of the mandible was limited. Hemihypoplasia also exerted an influence on structures that were not of branchiogenic origin, e.g., the cranial base (narrowing, asymmetry, and more pronounced curvature), the neurocranium (depression in mastoid and tympanotemporal regions, posterior rotation of the vault), and the frontonasal component (deviation of the nose and premaxilla). The cranial vault and the bottom of the occipital bone showed on the average no asymmetries. The similar character of deviations in slightly affected groups revealed that in spite of the high variability of changes typical for branchiogenic malformations the development of the face in these defects was subjected to certain rules. Marked facial asymmetry occurred only in every fifth patient with a severe degree of microtia, while definite signs of asymmetry were absent in every third patient.     

Effects of fronto-occipital cranial reshaping on mandibular form.

Cultural reshaping (artificial deformation or modification) of the neurocranial vault provides an artificially increased range of morphological variability within which the relationship between the growing neurocranium and face can be investigated. We analyze crania which have been fronto-occipitally compressed to ascertain possible morphological effects on the mandible. We collected measures of mandibular breadth, length, and height from 82 modified (N = 48) and unmodified (N = 34) crania from a Peruvian Ancon series. Angle classification was also scored in order to investigate whether or not occlusal relationships were affected by neurocranial reshaping. Only intercondylar distance (posterior mandibular breadth) exhibited significant differences between unmodified and modified groups, though this difference was relatively small compared with vault deformation. The modified crania had a higher frequency of normal occlusion (Class I) than the unmodified crania. Increased intercondylar breadth in modified skulls is due to a cascade of effects which begin with a direct effect of the fronto-occipital deforming device on neurocranial shape (increased neurocranial width). The increase in mandibular breadth may be a compensatory response to increased cranial base breadth and maintains articulation between the cranial base and mandible. The increased posterior breadth, coupled with a slight decrease in mandibular depth, may contribute to the change in occlusal relationships suggested for this sample.     

Basicranial influence on overall cranial shape

This study examines the extent to which the major dimensions of the cranial base (maximum length, maximum breadth, and flexion) interact with brain volume to influence major proportions of the neurocranium and face. A model is presented for developmental interactions that occur during ontogeny between the brain and the cranial base and neurocranium, and between the neurobasicranial complex (NBC) and the face. The model is tested using exocranial and radiographic measurements of adult crania sampled from five geographically and craniometrically diverse populations. The results indicate that while variations in the breadth, length and flexion of the cranial base are mutually independent, only the maximum breadth of the cranial base (POB) has significant effects on overall cranial proportions, largely through its interactions with brain volume which influence NBC breadth. These interactions also have a slight influence on facial shape because NBC width constrains facial width, and because narrow-faced individuals tend to have antero-posteriorly longer faces relative to facial breadth than wide-faced individuals. Finally, the model highlights how integration between the cranial base and the brain may help to account for the developmental basis of some morphological variations such as occipital bunning. Among modern humans, the degree of posterior projection of the occipital bone appears to be a consequence of having a large brain on a relatively narrow cranial base. Occipital buns in Neanderthals, who have wide cranial bases relative to endocranial volume, may not be entirely homologous with the morphology occasionally evident in Homo sapiens.     

The relationship between cranial base and maxillo-facial morphology in Egyptian children

The aim of this study was to investigate the relationship between the cranial base and maxillo-facial morphology in Egyptian children. Data were obtained from 95 lateral cephalograms for 61 boys and 34 girls aged from 7.5 to 9.5 years with mean age 8.5 years. Eighteen linear and 14 angular measurements were derived from 40 landmarks and recorded from the standardized radiographs that were traced following methods formerly defined and described. A principal component analysis of linear and angular measurements showed that anterior and posterior cranial base lengths and cranial base angle were closely associated in different ways with different aspects of maxillo-facial morphology in both sexes. This was more pronounced in boys. A significant positive relationship was found between anterior cranial base length and most of the variables describing the maxillo-facial morphology in both sexes. Posterior cranial base length was significantly correlated to the facial depth. The cranial base angle showed a significant negative correlation with the antero-posterior position of maxilla and mandible (SNA=-0.34, SNB=-0.27 and ANB=-0.24). In conclusion, cranial base configuration plays an important role in maxillo-facial morphology.     

Morphological evolution through integration: a quantitative study of cranial integration in Homo, Pan, Gorilla and Pongo

Morphological integration refers to coordinated variation among traits that are closely related in development and/or function. Patterns of integration can offer important insight into the structural relationship between phenotypic units, providing a framework to address questions about phenotypic evolvability and constraints. Integrative features of the primate cranium have recently become a popular subject of study. However, an important question that still remains under-investigated is: what is the pattern of cranial shape integration among closely related hominoids? To address this question, we conducted a Procrustes-based geometric morphometrics study to quantify and analyze shape covariation patterns between different cranial regions in Homo, Pan, Gorilla and Pongo. A total of fifty-six 3D landmarks were collected on 407 adult individuals. We then sub-divided the landmarks corresponding to cranial units as outlined in the ‘functional matrix hypothesis.’ Sub-dividing the cranium in this manner allowed us to explore patterns of covariation between the face, basicranium and cranial vault, using the two-block partial least squares approach. Our results suggest that integrated shape changes in the hominoid cranium are complex, but that the overall pattern of integration is similar among human and non-human apes. Thus, despite having very distinct morphologies the way in which the face, basicranium and cranial vault covary is shared among these taxa. These results imply that the pattern of cranial integration among hominoids is conserved.     

Craniofacial suture stenosis: morphologic effects

Craniofacial anomalies, such as Apert's and Crouzon's syndromes, are presumed to be related to premature growth arrest of cranial base growth sites. However, premature growth arrest at cranial vault sutures in animals appears to play a causative role in the development of cranial deformities characteristic of single-suture, or simple, craniosynostosis in humans. To study the possible causative role of cranial vault and other (interface) suture stenoses on the development of craniofacial deformity, a vault suture and an interface suture between the cranial vault and facial skeleton were simultaneously immobilized. Thirty-one New Zealand White rabbits at 9 days of age underwent implantation of dental amalgam growth markers adjacent to cranial vault and facial sutures. In the experimental group (n = 15), methylcyanoacrylate adhesive was applied over the coronal (vault) and frontonasal (interface suture between vault and facial skeleton) sutures to immobilize them. The remaining 16 animals served as sham-treated controls. All animals underwent serial radiographic cephalometry to document growth effects in the cranial vault, cranial base, and facial skeleton. Application of adhesive resulted in statistically significant (p less than 0.05) reduction in growth at the coronal and frontonasal sutures. This was accompanied by an overall significant reduction in neurocranial vault length during the first 30 days of development.(ABSTRACT TRUNCATED AT 250 WORDS)     

Evaluation of hand asymmetry in relation to hand preference

We evaluated the asymmetric hand measurements in right- and left-handed individuals. 343 men and 290 women aged 18-42 years (22.11 +/- 2.07) participated in the study. There were no statistically significant differences when right-left differences in hand length, third finger length, palmar length, and the digit index value were evaluated according to hand preference and sex. Statistically significant differences were found for right-left differences in hand width, hand-shape index, and the palmar length/width according to hand preference. The strong left-handers, weak left-handers, and ambidextrous individuals in the study group all exhibited asymmetry favoring the left and were considered together. Similarly, the strong and weak right-handers exhibited asymmetry favoring the right hand and were considered together. The difference between these two groups was significant. When the data were evaluated according to sex, significant differences were found between the subgroups. In particular, right-left differences in the hand-shape index and palmar length/width values of the strong left-handers, weak left-handers, and ambidextrous individuals were found to be statistically significant according to sex; in contrast, the strong and weak right-handers showed no significant differences according to sex. These results suggest a relation of hand asymmetry to hand preference in a Turkish population.     

Cranial integration in Homo: singular warps analysis of the midsagittal plane in ontogeny and evolution

This study addresses some enduring issues of ontogenetic and evolutionary integration in the form of the hominid cranium. Our sample consists of 38 crania: 20 modern adult Homo sapiens, 14 sub-adult H. sapiens, and four archaic Homo. All specimens were CT-scanned except for two infant H. sapiens, who were imaged by MR instead. For each specimen 84 landmarks and semi-landmarks were located on the midsagittal plane and converted to Procrustes shape coordinates. Integration was quantified by the method of singular warps, a new geometric-statistical approach to visualizing correlations among regions. The two classic patterns of integration, evolutionary and ontogenetic, were jointly explored by comparing analyses of overlapping subsamples that span ranges of different hypothetical factors. Evolutionary integration is expressed in the subsample of 24 adult Homo, and ontogenetic integration in the subsample of 34 H. sapiens. In this data set, vault, cranial base, and face show striking and localized patterns of covariation over ontogeny, similar but not identical to the patterns seen over evolution. The principal differences between ontogeny and phylogeny pertain to the cranial base. There is also a component of cranial length to height ratio not reducible to either process. Our methodology allows a separation of these independent processes (and their impact on cranial shape) that conventional methods have not found.     

Craniofacial changes in unilateral microtia: II. An X-ray study

Roentgenocephalometry was used for studies into the extent and character of craniofacial changes in 45 adult males with unilateral (right-sided) microtia. Out of the whole complex of changes associated with this malformation the mandibular ramus showed the most marked involvement and represented the main cause of the accompanying deviations and asymmetries. On the average, the affected half of the face was compressed toward the level of the external auditory meatus both from above and below, but there was a marked variability in individual patients examined. No signs of asymmetry were disclosed in one third of the patients while severe asymmetry was present in one fifth of the patients. Facial hemihypoplasia exerted no substantial influence on the facial profile (when no retrusion of the lower jaw was present) on the sagittal maxillomandibular relations or on the occlusion of incisors, while in transverse direction a laterosuperior deviation of the mandible towards the affected side was clearly visible. A branchiogenic malformation affected the neighboring structures, the cranial base (a more marked curving), frontonasal segment (septum and premaxillar deviation), and the neurocranium (posterior rotation of the cranial vault). The inner ear structures (semicircular canals) were affected only rarely (in 4% of patients). These findings complemented the results obtained in the first part of our study and confirmed the complex character of this inborn anomaly.     

Directional asymmetry in the limbs,skull and pelvis of the silver fox (V. vulpes)

Directional asymmetry (DA) is a characteristic of most vertebrates, most strikingly exhibited by the placement of various organs (heart, lungs, liver, etc.) but also noted in small differences in the metrics of skeletal structures such as the pelvis of certain fish or sauropsids. We have analyzed DA in the skeleton of the fox (V. vulpes), using ～1,000 radiographs of foxes from populations used in the genetic analysis of behavior and morphology. Careful measurements from this robust data base demonstrate that: 1) DA occurs in the limb bones, the ileum, and ischium and in the mandible; 2) regardless of the direction of the length asymmetry vector of a particular skeletal unit, the vectorial direction of length is always opposite to that of width; 3) with the exception of the humerus and radius, there is no correlation or inverse correlation between vectorial amplitudes or magnitudes of bone asymmetries. 4) Postnatal measurements on foxes demonstrate that the asymmetry increases after birth and continues to change (increasing or decreasing) during postnatal growth. 5) A behavior test for preferential use of a specific forelimb exhibited fluctuating asymmetry but not DA. None of the skeletal asymmetries were significantly correlated with a preferential use of a specific forelimb. We suggest that for the majority of fox skeletal parameters, growth on the right and left side of the fox are differentially biased resulting in fixed differences between the two sides in either the rate of growth or the length of the period during which growth occurs. Random effects around these fixed differences perturb the magnitude of the effects such that the magnitudes of length and width asymmetries are not inversely correlated at the level of individual animals. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

Three-dimensional geometric morphometric analysis of cranio-facial sexual dimorphism in a Central European sample of known sex

This article presents an approach for estimating the sexual dimorphism of adult crania using three-dimensional geometric morphometric methods. The study sample consisted of 139 crania of known sex (73 males and 66 females) belonging to persons who lived during the first half of the 20th century in Bohemia. The three-dimensional co-ordinates of 82 ecto-cranial landmarks and 39 semi-landmarks covering the midsagittal curve of the cranial vault were digitised using a MicroScribe G2X contact digitiser. The purposes of the investigation were to define the regions of the cranium where sexual dimorphism is most pronounced and to investigate the effectiveness of this method for determining sex from the shape of the cranium. The results demonstrate that it is better to analyse apportionable parts of the cranium rather than the cranium as a whole. Significant sexual differences (significance was determined using multivariate analysis of variance) were noted in the shape of the midsagittal curve of the vault, upper face, the region of the nose, orbits, and palate. No differences were recorded either in the shape of the cranium as a whole or in the regions of the base and the neurocranium. The greatest accuracy in determining sex was found in the region of the upper face (100% of study subjects correctly classified) and the midsagittal curve of the vault (99% of study subjects correctly classified).     

Sexual size dimorphism and allometric growth of Morelet's crocodiles in captivity

Few studies have conducted morphological analyses of crocodilians, and little information exists on differences between size-classes and sexes in Neotropical crocodilians. In this study, we measured nine morphological traits in 121 captive Morelet's crocodiles Crocodylus moreletii (81 females and 40 males). Our results revealed that individuals < 2 m total length do not exhibit sexual dimorphism in morphometric characteristics. However, for crocodiles over 2 m in length, males were significantly larger than females in terms of dorsal-cranial length, cranial width, snout width and snout-ventral length. In general, morphological traits demonstrated a strongly significant relationship with total length at the smaller size class of 150-200 cm length. However, in the highest size class of 250-300 cm length (large adult males), morphological traits were no longer significantly related with total length. Male crocodiles demonstrated allometric growth of cranial morphology with significantly greater increase in cranial width, snout width, and mid-snout width relative to total length at higher size classes. Morphological dimorphism and allometric growth may be associated with adaptive strategies for reproductive success.     

A factor analysis of cranial base and vault dimensions in children

Lengths within the cranial base and vault were measured in cephalometric radiographs of 220 boys and 177 girls ranging in age from 0 to 15 years; all these children are participants in The Fels Longitudinal Growth Study. The present study is based on mixed longitudinal data derived from 1640 radiographs for boys and 1260 radiographs for girls. Factor analysis was applied separately for boys and girls for each age group; i.e., 0–3, 4–6, 7–9, 10–12, and 13–15 years. For the 0–3 year age group, two factors were extracted in each sex, whereas four factors were extracted in the rest of the age groups. The factor structures are similar in the three older age groups of boys (7–9, 10–12, and 13–15 years). The first four factors for these groups are labelled, respectively: cranial vault size, posterior cranial base length, presphenoid length, and basisphenoid length. The order of the third and fourth factors is reversed in the 7–9 year olds. For girls, the factors extracted were also the same in both the 7–9 and 10–12 year age groups, even though the order of factors was different between age groups; i.e., anterior cranial base length, cranial vault size, basisphenoid length, and basioccipital length. Differential growth rates among cranial base dimensions probably cause changes in factor patterns. Obliteration of the spheno-occipital synchondrosis is suggested as the mechanism responsible for the change of factor pattern in the girls. Closure of this synchondrosis would have occurred too late to affect the patterns in boys.     

Temporal squama shape in fossil hominins: relationships to cranial shape and a determination of character polarity

In 1943, Weidenreich described the squamosal suture of Homo erectus as long, low, and simian in character and suggested that this morphology was dependent upon the correlation between the size of the calvarium and the face. Many researchers now consider this character to be diagnostic of H. erectus. The relationship between cranial size and shape and temporal squama morphology, however, is unclear, and several authors have called for detailed measurements of squamosal variation to be collected before any conclusions are drawn regarding the nature of the morphology observed in H. erectus. Thirteen fossil and extant taxa were examined to address two questions: 1) Are size and shape of the temporal squama correlated with cranial vault morphology? and 2) Is the H. erectus condition plesiomorphic? To answer these questions, measurements were collected and indices were calculated for squamosal suture height, length, and area in relation to metric variables describing cranial size and shape. A two‐dimensional morphometric study was also completed using High Resolution‐Polynomial Curve Fitting (HR‐PCF) to investigate correlations between curvature of the squamosal suture and curvature of the cranial vault. Results of both analyses indicate that squamosal suture form is related to cranial size and shape. Furthermore, the plesiomorphic condition of the squamosal suture for hominins was identified as high and moderately arched; this condition is retained in H. erectus and is distinct from the great ape condition. It is suggested that this similarity is the result of increased cranial length without a corresponding increase in cranial height. Am J Phys Anthropol, 2008. © 2008 Wiley‐Liss, Inc.     

Patterns and development of floral asymmetry in Senna (Leguminosae, Cassiinae)

The buzz-pollinated genus Senna (Leguminosae) is outstanding for including species with monosymmetric flowers and species with diverse asymmetric, enantiomorphic (enantiostylous) flowers. To recognize patterns of homology, we dissected the floral symmetry character complex and explored corolla morphology in 60 Senna species and studied floral development of four enantiomorphic species. The asymmetry morph of a flower is correlated with the direction of spiral calyx aestivation. We recognized five patterns of floral asymmetry, resulting from different combinations of six structural elements: deflection of the carpel, deflection of the median abaxial stamen, deflection or modification in size of one lateral abaxial stamen, and modification in shape and size of one or both lower petals. Prominent corolla asymmetry begins in the earl-stage bud (unequal development of lower petals). Androecium asymmetry begins either in the midstage bud (unequal development of thecae in median abaxial stamen; twisting of androecium) or at anthesis (stamen deflection). Gynoecium asymmetry begins in early bud (primordium off the median plane, ventral slit laterally oriented) or midstage to late bud (carpel deflection). In enantiostylous flowers, pronouncedly concave and robust petals of both monosymmetric and asymmetric corollas likely function to ricochet and direct pollen flow during buzz pollination. Occurrence of particular combinations of structural elements of floral symmetry in the subclades is shown.