Ontogenic growth changes of the skull base in four genera of nonhuman primates.

Cross-sectional studies of the degree of the cranial base flexion were carried out in infant, juvenile and adult skulls in four genera of nonhuman primates (P. paniscus, H. lar, P. urinus, and M. mullatta). The cephalometric observations of the cranial base included linear and angular measurements of each specimen. The data obtained in this study showed that the anterior portion of the cranial base exhibits a significant shortening trend as the mammalian evolutionary scale ascends. Moreover, the growth pattern of the anterior portion of the skull base follows that of the facial bony structures. The ontogenic growth changes of the posterior portion of the skull base follows the growth pattern of the endocranial cavity. The significant trend of elongation in this area directly contributes to the posterior migration of the foramen magnum. The magnitude of these growth changes decreases as the evolutionary scale ascends. The angular measurements of the cranial flexion showed a less obtuse cranial base angle in young specimens and the ones higher on the mammalian scale. The skull kyphosis was less pronounced in these specimens and the anatomical features of the cranial base were more humanlike, including the balance of the head expressed by the position of the foramen magnum.     

Postnatal growth of skull linear measurements of Cape Hare Lepus capensis in northern China: an analysis in an adaptive context

304 skulls of Cape hare (Lepus capensis) were collected from two climatically distinct localities in northern China. With eye lens weight as a continuous age variable, postnatal growth patterns of 25 cranial linear measurements in relation to sex, growth season and region were analysed to understand the morphological basis of life history adaptation. In almost all the skull measurements, no significant differences were found between either sex or growth seasons. Principal component analysis revealed that facial elements accounted for the greatest proportion of skull morphological variation. Von Bertalanffy function was applied to describe growth trajectories of the skull elements. Based on this model, the growth rates of skull elements and the age at which they reached a certain proportion (95%) of asymptotes were compared. The results showed that skull growth exhibited an allometric pattern, with neural components attaining their final size more rapidly (at about 2–3 months old in tympanic bulla and 4–6 months old in others) than did the facial, which continued to grow well into postnatal life (at 6–10 months old). The earlier establishment of neurocranial morphology was associated with a fully developed central nervous system, which may play a key role in improving the survival of animals during the early phase of life. There was a regional difference in developmental rate of the hare skull. For all the skull parameters, northern hares had a more rapid rate of cranial growth compared to the southern, i.e. skull elements of juveniles from northern population were relatively larger at comparable ages and achieved adult size 0.5–4.0 months earlier than those from the south. In adult hares, however, no significant regional differences in any of the skull parameters were present. Adaptive explanations for the regional difference in ontogenetic pattern of skull morphology include age‐specific thermoregulation constraint, season‐related food availability and age‐dependent predation pressure. Based on the findings of this study, it is suggested that the postnatal growing period represents a crucial time of life, and that improvement of survivorship when young by growth adaptation forms an important aspect of the hare's life history strategies. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 78 , 343–353.     

Avian skull morphological evolution: exploring exo- and endocranial covariation with two-block partial least squares

While rostral variation has been the subject of detailed avian evolutionary research, avian skull organization, characterized by a flexed or extended appearance of the skull, has eventually become neglected by mainstream evolutionary inquiries. This study aims to recapture its significance, evaluating possible functional, phylogenetic and developmental factors that may be underlying it. In order to estimate which, and how, elements of the skull intervene in patterning the skull we tested the statistical interplay between a series of old mid-sagittal angular measurements (mostly endocranial) in combination with newly obtained skull metrics based on landmark superimposition methods (exclusively exocranial shape), by means of the statistic-morphometric technique of two-block partial least squares. As classic literature anticipated, we found that the external appearance of the skull corresponds to the way in which the plane of the caudal cranial base is oriented, in connection with the orientations of the plane of the foramen magnum and of the lateral semicircular canal. The pattern of covariation found between metrics conveys flexed or extended appearances of the skull implicitly within a single and statistically significant dimension of covariation. Marked shape changes with which angles covary concentrate at the supraoccipital bone, the cranial base and the antorbital window, whereas the plane measuring the orientation of the anterior portion of the rostrum does not intervene. Statistical covariance between elements of the caudal cranial base and the occiput inplies that morphological integration underlies avian skull macroevolutionary organization as a by-product of the regional concordance of such correlated elements within the early embryonic chordal domain of mesodermic origin.     

The sixth skull cap of Pithecanthropus erectus

The sixth skull cap of Pithecanthropus erectus (or skull V, since the Modjokerto skull has not been given a number) was found in the upper layers of the Trinil beds of Sangiran (Central Java) in 1963, associated with fossils of the Sino-Malayan fauna. No stone tools were discovered in direct association with the find. The specimen consists of the occipital, both parietals, both temporals, sphenoid fragments, the frontal and the left zygomatic bone. We consider the skull to be a male in his early twenties. The occipital, parietal, frontal and temporal bones demonstrate definite pithecanthropine characteristics, and the cranial capacity is estimated to be 975 cm³. Of the superstructures, the supraorbital torus is extraodinarily thick, approaching the condition in Australopithecus boisei and Rhodesian man. And the sagittal torus is certainly higher than in skulls I and II, but lower than in skull IV. In addition, the angle between the occipital and nuchal planes is larger than in the previous finds. As revealed by various features, the gap between the robustness of skull IV on one hand, and skulls I, II and III on the other, is bridged by the present find. There is no reasonable taxonomic need to ascribe this specimen to a new species, because it seems to be merely an intrapopulational variant of the same species. Other skulls of P. erectus suggest that the bregmatic eminence, and hence the vertex, is invariably situated at bregma, but this new skull cap deviates from the pattern. Its pteric regions disclose the anthropoid X and I types. The middle meningeal groove pattern is similar to other Pithecanthropus skulls; however, it betrays a known anomaly in that the main stem is covered for a short distance by a bony plate. The mastoid process is fairly well developed, and is also well pneumatized as in P. pekinensis, with its air cells invading the pronounced supramastoid crest. The zygomatic bone, the first one recovered of P. erectus, does not show characters of particular importance. In fact, its thickness is in the range of modern man. We would like to stress that the absence of the cranial base does not necessarily indicate that the specimen must be a poor victim of cannibalism, since the morphology of the base renders it more susceptible to post-mortem natural traumata.     

Infant skull and suture properties: measurements and implications for mechanisms of pediatric brain injury

The mechanical properties of the adult human skull are well documented, but little information is available for the infant skull. To determine the age-dependent changes in skull properties, we tested human and porcine infant cranial bone in three-point bending. The measurement of elastic modulus in the human and porcine infant cranial bone agrees with and extends previous published data [McPherson, G. K., and Kriewall, T. J. (1980), J. Biomech., 13, pp. 9-16] for human infant cranial bone. After confirming that the porcine and human cranial bone properties were comparable, additional tensile and three-point bending studies were conducted on porcine cranial bone and suture. Comparisons of the porcine infant data with previously published adult human data demonstrate that the elastic modulus, ultimate stress, and energy absorbed to failure increase, and the ultimate strain decreases with age for cranial bone. Likewise, we conclude that the elastic modulus, ultimate stress, and energy absorbed to failure increase with age for sutures. We constructed two finite element models of an idealized one-month old infant head, one with pediatric and the other adult skull properties, and subjected them to impact loading to investigate the contribution of the cranial bone properties on the intracranial tissue deformation pattern. The computational simulations demonstrate that the comparatively compliant skull and membranous suture properties of the infant brain case are associated with large cranial shape changes, and a more diffuse pattern of brain distortion than when the skull takes on adult properties. These studies are a fundamental initial step in predicting the unique mechanical response of the pediatric skull to traumatic loads associated with head injury and, thus, for defining head injury thresholds for children.     

Basicranial axis length v. skull length in analysis of carnivore skull shape

Skull length is the measurement most commonly used as a standard against which other aspects of cranial morphology are compared to derive an index of relative size or proportions. However, skull length is composed of two different functional components, facial skull and cerebral skull, which vary independently and have different scaling relationships with body size. An analysis of carnivore skull shape with measurements standardized against basicranium length produced very different results than an analysis using skull length as the standard. For example, expressions of relative size of cranial measurements were reduced by 13% in mustelids and increased by 20% in canids, reflecting removal of jaw length (short in mustelids and long in canids) from the comparative standard (basicranial axis length). Cranial measurements scale with higher allometric exponents against basicranial axis length than against skull length.     

Measurements on the grade of the human skull base and angle determinations]

1. The angle between the Os sphenoidale and the Clivus has been determined on 106 skulls. The mean was 117.68 degrees. 2. The altitude of the Foramen magnum with respect to the Frankfort plane was determined at -1.96 cm on the average. 3. The angle of inclination of the Foramen magnum to the Frankfort plane was determined from the altitude of the Basion and Opisthion and from the length of the Foramen magnum. The mean was -13.71 degrees. 4. In the inner base of the skull the altitudes of following structures have been determined: Planum sphenoidale (mean 2.34 cm), ground of the Fossa hypophysialis (1.32 cm), ground of the middle cranial fossa (-0.03 cm), ground of the posterior cranial fossa (-2.30 cm), upper edge of the Os petrosum (1.12 cm), Tuberculum jugulare (-0.21 cm), ground of the Meatus acusticus internus (0.45 cm). 5. In order to provide numerical data for the step-like building of the inner base of the skull, the differences between the altitudes of important structures from the inner base have been determined: Margo superior partis petrosae - Tuberculum jugulare (mean 1.35 cm). Opisthion - ground of the posterior cranial fossa (0.11 cm), Planum sphenoidale - ground of the middle cranial fossa (2.38 cm), ground of the middle cranial fossa - ground of the posterior cranial fossa (2.27 cm), Margo superior partis petrosae - ground of the posterior cranial fossa (3.35 cm), Margo superior partis petrosae - ground of the middle cranial fossa (1.15 cm).     

Height of skull base as an indicator of living conditions in historical native populations from Europe, Australia and Africa

Flattening and lowering of the skull base in response to improper bone growth is called platybasis, and is considered a sensitive and reliable indicator of adverse living conditions such as malnutrition and disease during the prenatal period and early childhood. The degree of platybasis was assessed in three series of skulls representing geographically distinct historical native populations from Europe, Australia and Africa. Platybasis was determined by measuring the height of the base of the skull. The degree of platybasis varied among the populations examined, and was the lowest in the Australian group, and the highest in the African group. This may be due to either variability of living conditions or genetic factors, which have an influence on robusticity and cranial architecture. There were also differences among the groups in terms of the other skull measurements and indices examined. The height of the base of the skull was generally greater in males than in females, which indicates sexual dimorphism or fact that females had worse living conditions. Correlation coefficients between the height of the base of the skull and other measurements including skull length and skull width were also calculated indicating significant relationships. The differences between the height of the base of the skull and height/length index and height/width index, were statistically significant.     

Influence of the lateral ventricles and irregular skull base on brain kinematics due to sagittal plane head rotation

Two-dimensional physical models of the human head were used to investigate how the lateral ventricles and irregular skull base influence kinematics in the medial brain during sagittal angular head dynamics. Silicone gel simulated the brain and was separatedfrom the surrounding skull vessel by paraffin that provided a slip interface between the gel and vessel. A humanlike skull base model (HSB) included a surrogate skull base mimicking the irregular geometry of the human. An HSBV model added an elliptical inclusion filled with liquid paraffin simulating the lateral ventricles to the HSB model. A simplified skull base model (SSBV) included ventricle substitute but approximated the anterior and middle cranial fossae by a flat and slightly angled surface. The models were exposed to 7600 rad/s2 peak angular acceleration with 6 ms pulse duration and 5 deg forced rotation. After 90 deg free rotation, the models were decelerated during 30 ms. Rigid body displacement, shear strain and principal strains were determined from high-speed video recorded trajectories of grid markers in the surrogate brains. Peak values of inferior brain surface displacement and strains were up to 10.9X (times) and 3.3X higher in SSBV than in HSBV. Peak strain was up to 2.7X higher in HSB than in HSBV. The results indicate that the irregular skull base protects nerves and vessels passing through the cranial floor by reducing brain displacement and that the intraventricular cerebrospinal fluid relieves strain in regions inferior and superior to the ventricles. The ventricles and irregular skull base are necessary in modeling head impact and understanding brain injury mechanisms.     

Use of the free vastus lateralis flap in skull base reconstruction

Free flaps in skull base reconstruction are indicated for providing an effective separation of the intracranial cavity from the oronasal space, for eliminating a dead space, and for the treatment of established wound complications such as dural exposures and cerebrospinal fluid leaks. Seven patients with cranial base defects underwent reconstructions using a free vastus lateralis muscle flap. In two cases, a vastus lateralis flap was raised to incorporate the anterolateral thigh skin as a myocutaneous flap. In four cases, a free flap was indicated for reconstruction following tumor ablation, and in three cases, for the resolution of wound or cerebrospinal fluid leak complications following previous cranial base surgery. All flaps were successful, with no partial failures. In those patients undergoing tumor ablative surgery, the cranial cavity was effectively sealed from the oronasal cavity. Patients with established wound complications following previous cranial base surgery had a complete resolution of their symptoms. This report discusses the suitability of the vastus lateralis flap for skull base reconstruction in terms of the availability of adequate muscle volume to fill dead space, vascularized fascia to augment dural repairs, and the freedom to use skin if required for internal lining or external skin cover. This flap also provides an extremely long pedicle, allows simultaneous flap harvest, and has low donor site morbidity.     

Patterns of morphological integration between parietal and temporal areas in the human skull

Modern humans have evolved bulging parietal areas and large, projecting temporal lobes. Both changes, largely due to a longitudinal expansion of these cranial and cerebral elements, were hypothesized to be the result of brain evolution and cognitive variations. Nonetheless, the independence of these two morphological characters has not been evaluated. Because of structural and functional integration among cranial elements, changes in the position of the temporal poles can be a secondary consequence of parietal bulging and reorientation of the head axis. In this study, we use geometric morphometrics to test the correlation between parietal shape and the morphology of the endocranial base in a sample of adult modern humans. Our results suggest that parietal proportions show no correlation with the relative position of the temporal poles within the spatial organization of the endocranial base. The vault and endocranial base are likely to be involved in distinct morphogenetic processes, with scarce or no integration between these two districts. Therefore, the current evidence rejects the hypothesis of reciprocal morphological influences between parietal and temporal morphology, suggesting that evolutionary spatial changes in these two areas may have been independent. However, parietal bulging exerts a visible effect on the rotation of the cranial base, influencing head position and orientation. This change can have had a major relevance in the reorganization of the head functional axis.     

Pervasive genetic integration directs the evolution of human skull shape

It has long been unclear whether the different derived cranial traits of modern humans evolved independently in response to separate selection pressures or whether they resulted from the inherent morphological integration throughout the skull. In a novel approach to this issue, we combine evolutionary quantitative genetics and geometric morphometrics to analyze genetic and phenotypic integration in human skull shape. We measured human skulls in the ossuary of Hallstatt (Austria), which offer a unique opportunity because they are associated with genealogical data. Our results indicate pronounced covariation of traits throughout the skull. Separate simulations of selection for localized shape changes corresponding to some of the principal derived characters of modern human skulls produced outcomes that were similar to each other and involved a joint response in all of these traits. The data for both genetic and phenotypic shape variation were not consistent with the hypothesis that the face, cranial base, and cranial vault are completely independent modules but relatively strongly integrated structures. These results indicate pervasive integration in the human skull and suggest a reinterpretation of the selective scenario for human evolution where the origin of any one of the derived characters may have facilitated the evolution of the others.     

Morphometric variation of the skull during postnatal development in the Lowland European bisonBison bonasus bonasus

We studied the variation of linear measurements and skull capacity in Lowland European bisonBison bonasus bonasus (Linnaeus, 1758) during postnatal development, and the dependencies of the parameters in relation to sex, age, and body mass of the animals. Material consisted of 599 bison skulls (310 males and 289 females), within the age range of 1 month to 21 years (males) and to 27 years (females). In the group of calves to 1 year old, no sex connected differences in skull measurements were observed, whereas the skull capacity in older calves was significantly larger (0.01>p>0.001) in males than in females. From the third year of life, most skull measurements display characteristics of sexual dimorphism. Skull development in both sexes is most intensive during the first three years of life, and slows from the age of 5. In older individuals of both sexes (≥ 6 years), orbital breadth continues growing and, in females, breadth of splanchnocranium continues increasing. Growth in a bison’s skull capacity is most intensive up to the third year of life and slows from the age of 5. During postnatal development, a bison skull grows proportionally except the neurocranium, which grows slightly slower in comparison with basal length and its development finishes earlier than that of splanchnocranium. In ontogenesis, a bison skull grows much slower compared to body mass. In relation to body mass, skull capacity and the height of neurocranium grow most slowly while orbital breadth grows most intensively. The results obtained were compared with data on skull sizes of bison born in 1930–1950 and bred in captivity and with skulls of the American bisonBison bison. Inbreeding is probably responsible for some types of phenotypic abnormalities in the skull which appear in modern European bison.     

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.     

Dispelling dog dogma: an investigation of heterochrony in dogs using 3D geometric morphometric analysis of skull shape

Heterochrony is an evolutionary mechanism that generates diversity via perturbations of the rate or timing of development that requires very little genetic innovation. As such, heterochrony is thought to be a common evolutionary mechanism in the generation of diversity. Previous research has suggested that dogs evolved via heterochrony and are paedomorphic wolves. This study uses three-dimensional landmark-based coordinate data to investigate heterochronic patterns within the skull morphology of the domestic dog. A total of 677 adult dogs representing 106 different breeds were measured and compared with an ontogenetic series of 401 wolves. Geometric morphometric analysis reveals that the cranial shape of none of the modern breeds of dogs resembles the cranial shapes of adult or juvenile wolves. In addition, investigations of regional heterochrony in the face and neurocranium also reject the hypothesis of heterochrony. Throughout wolf cranial development the position of the face and the neurocranium remain in the same plane. Dogs, however, have a de novo cranial flexion in which the palate is tilted dorsally in brachycephalic and mesaticephalic breeds or tilted ventrally in dolichocephalic and down-face breeds. Dogs have evolved very rapidly into an incredibly morphologically diverse species with very little genetic variation. However, the genetic alterations to dog cranial development that have produced this vast range of phylogenetically novel skull shapes do not coincide with the expectations of the heterochronic model. Dogs are not paedomorphic wolves.     

Studies on orthocephalization. 2. Flexions of the rat head in the period between 14 and 60 days after gestation

The present paper considers the significance of a variety of cranial flexions in the process of orthocephalization of the rat skull between 14 and 60 days postnatally. The study is based on a sample of 27 male rats, who have been x-rayed at 14, 30 and 60 days with subsequent analyses of the photographs obtained. In this period the angle between the cranial base and the facial part of the skull becomes more obtuse, i.e. the skull becomes more orthocranial. The cranial base becomes at the same time more airobasal (lordotic). Angular changes between the individual bones in the cranial vault straightens the vault markedly. By this it becomes orthodorsal. As the angle between the basisphenoid and the parietal bones stays more or less constant between 14 and 60 days, the impression is created that both the anterior and posterior parts of the neural skull rotate upwards relative to this bone complex. Thus, orthocephalization in the period between 14 and 60 days both consists of flexions between the facial and neural parts of the skull (prebasal flexions), and may be more importantly of interosseous ventral and dorsal flexions.     

Cranial base and jaw relationship

The lateral skull radiographs of 124 boys aged approximately 10 years divided equally between the four angle classes were digitized in an effort to establish the relationship between cranial base size and shape and jaw relationship. Comparison of the means for occlusal groups showed a trend from class II to class III as cranial base dimensions and angle decreased. The condyle was also more distally positioned with respect to nasion, point A and the Pterygomaxillary vertical in the class II groups. Cranial base length correlated strongly with maxillary length but weakly with mandibular length. Nevertheless, the size of the maxilla did not influence its prognathism. The cranial base angle was strongly correlated (-0.7) with angle sella-nasion-point B. It is concluded that cranial base size and shape influence mandibular prognathism by determining the anteroposterior position of the condyle relative to the facial profile.     

Craniofacial deformity in patients with uncorrected congenital muscular torticollis: an assessment from three-dimensional computed tomography imaging

Congenital muscular torticollis is caused by idiopathic fibrosis of the sternocleidomastoid muscle that restricts movement and pulls the head toward the involved side. Deformation of the craniofacial skeleton will develop if the restriction is not released and result in aesthetic and functional problems. The purpose of this study was to use three-dimensional computed tomography imaging for qualitative and quantitative evaluation of the craniofacial deformity in a series of patients with uncorrected congenital muscular torticollis, and to assess age as a precipitating factor for severity of the deformity. A total of 14 patients from 1 month to 24 years of age were included. The skull images were rotated into standard orientation and reconfigured for evaluation of the cranium, endocranial base, and facial skeletal structures. The midlines of cranial base and facial bone, angle of midline deviation, width of each hemicranium and hemiface, and the orbital index were defined and measured. The results showed that the cranium and cranial base deformation took place as early as in infant stage, with the most prominent change occurring in the posterior cranial fossa. Facial bone asymmetry started to appear after 5 years of age, at which time the mandibular and occlusal abnormalities were observed. The deformity of the orbits and maxilla occurred at an older age, characterized by the deviation and decreased vertical height on the affected side. The severity of the observed deformities increased with age. The angle of midline deviation was 2.48 +/- 1.68 degrees in the cranial base and 3.26 +/- 3.28 degrees on the facial bone. Both of the midline deviations were significantly correlated with age. Compared with the contralateral side, the width of the ipsilateral posterior hemicranium was longer (54.36 +/- 6.72 mm versus 50.81 +/- 6.55 mm), and the width of the ipsilateral lower hemiface was shorter (35.30 +/- 7.27 mm versus 43.49 +/- 11.34 mm). Both differences were statistically significant. Measurement of the orbital index demonstrated a significantly flatter orbit on the ipsilateral side (89.48 +/- 0.11 versus 92.74 +/- 0.08). This study showed that the cranium and cranial base deformity occurred early in patients with uncorrected torticollis, while the facial bone deformity occurred in childhood stage. The cranial and facial deformity became more severe with age. Early release of the muscle restriction is advised to prevent craniofacial deformation.     

A small skull from Flores dated to the 20th century

A human skull with mandible from the Ngada District on the island of Flores, Indonesia, is described in order to contribute to the knowledge of variation in cranial architecture, which is important in interpretations of evolutionary cerebralisation. The skull was excavated in 1924 and sent to the National Museum in Copenhagen. The “Copenhagen Flores” (CF) male skull is radiocarbon-dated and of modern age. The cranium is small, but larger than e.g. Liang Bua skull (LB1) in every measurement. The (CT-scan based) cranial capacity of 1258 ml is normal for modern humans, but somewhat lower than values from the middle or upper Palaeolithics. The metric cranial data analysed in FORDISC, characterize the skull as a male Vietnamese rather than a Chinese or White individual. Tooth morphology shows the sundadont pattern and tooth size corresponds to that of teeth from Bali, Java and Malayan Orang Asli. Remarkable are the marked asymmetries in the dentition with rotation of an upper premolar and congenital absence of a third molar. In these respects the CF skull is similar to dentitions belonging to the pygmoid villagers of Rampasasa, a village not far from the Liang Bua cave, and to LB1.     

Diachronic variation in cranial thickness of Near Eastern populations

Cephalometric radiographs were taken of 111 skulls of skeletal remains of populations living in Israel and Jordan during the last 12,000 years. From these radiographs, skull length and height, and cranial thickness were measured. For each sex and period, high correlations were found between cranial thickness at vertex, bregma, and lambda. Cranial thickness at nasion was correlated with sinus width but not sinus height. All measurements were correlated with skull length but not skull breadth. Using multivariate analysis, no significant differences in cranial thickness were found between the sexes. Significant diachronic trends were found in lambda and sinus width, and they were independent of variation in skull length.