Surface facial modelling and allometry in relation to sexual dimorphism

Sexual dimorphism is responsible for a substantial part of human facial variability, the study of which is essential for many scientific fields ranging from evolution to special biomedical topics. Our aim was to analyse the relationship between size variability and shape facial variability of sexual traits in the young adult Central European population and to construct average surface models of adult males and females. The method of geometric morphometrics allowed not only the identification of dimorphic traits, but also the evaluation of static allometry and the visualisation of sexual facial differences.Facial variability in the studied sample was characterised by a strong relationship between facial size and shape of sexual dimorphic traits. Large size of face was associated with facial elongation and vice versa. Regarding shape sexual dimorphic traits, a wide, vaulted and high forehead in combination with a narrow and gracile lower face were typical for females. Variability in shape dimorphic traits was smaller in females compared to males. For female classification, shape sexual dimorphic traits are more important, while for males the stronger association is with face size.Males generally had a closer inter-orbital distance and a deeper position of the eyes in relation to the facial plane, a larger and wider straight nose and nostrils, and more massive lower face. Using pseudo-colour maps to provide a detailed schematic representation of the geometrical differences between the sexes, we attempted to clarify the reasons underlying the development of such differences.     

Investigation of mandibular and neurocranial form

In a study of mandibular form, 19 traditional linear dimensions were found to be consistently greater for macrocephalics than microcephalics. As such dimensions combine both size and shape parameters together, their interpretation proved difficult. In order to examine specifically mandibular shape only, each mandibular outline form was digitized and subjected to the technique of medial axis transformation. The data indicate that the shape contrasts between the mandibles of macrocephalics and microcephalics are more complex than traditionally envisaged and emphasize the need for further study to ascertain the effect of genetic and environmental influences on mandibular form.     

腊尔山区苗族学生头面部形态特征

本项目对1896例腊尔山区6~16岁苗族学生（男919例,女977例）进行了人体测量,测量了10项头面部形态指标,并根据公式计算13项头面部指数值。结果表明,苗族学生头面部各指标均值随年龄增长而增加,同年龄段头面部各指标均值男生一般高于女生,且多数年龄段差异有统计学意义;男性的头长宽的、形态面的、形态上面的、容貌面的、颧额宽的、容貌上面的和容貌上面高的指数以及女性的形态面的、形态上面的、容貌面的、颧下颌宽度的、容貌上面高的指数随年龄增长总体趋势是先增加后递减;男性头面高的以及女性的头长高的、头宽高的和头面宽的指数随年龄增长逐渐增加;男女性之间的头长高的、头宽高的和额顶宽的指数比较显示,多数年龄段差异有统计学意义;苗族学生头型以中头型、高头型、阔头型多见,面型以阔面型、阔上面型为主;相对于其他族群,腊尔山苗族男生面型短而宽,头型偏长,女生面型偏短,头型稍偏长而窄。腊尔山苗族学生头面部特征与融水苗族、土家族等族群较为接近,与哈萨克族、新乡汉族相距较远。     

Variation in soft-tissue thicknesses on the human face and their relation to craniometric dimensions

The average thickness of soft tissues on parts of the face is known, but its variation has not been related to cranial morphology. To investigate this relationship, measurements of facial soft-tissue depths and craniometric dimensions were taken on adult, white Australian cadavers (17 male and 23 female). Significant correlations between many soft-tissue depths and craniometric dimensions were found, suggesting a relationship between the amount of soft tissue present on the face and the size of the underlying bony skeleton. Soft-tissue depths were highly positively correlated with each other; craniometric dimensions were correlated but to a lesser extent. Males had thicker soft tissues and larger craniometric dimensions than females; considerable overlap of ranges was also noted. Multiple regression analysis was used to produce equations predicting the soft-tissue depth at specified areas of the face from craniometric dimensions. A subsample of nine cadavers was examined for the effects of tissue embalming. Embalming caused significant initial increases in facial soft-tissue depths. Cadavers embalmed for less than 6 months had soft-tissue depths significantly greater than for fully embalmed cadavers. The evidence that facial soft-tissue thicknesses vary with craniofacial dimensions has implications for forensic identification, facial aesthetic surgery, and approximation of the facial features of extinct individuals.     

Correlations between the cross-sectional area of the jaw muscles and craniofacial size and shape

In adult human subjects, the correlations were determined between the cross-sectional areas of the jaw muscles (measured in CT scans) and a number of facial angles and dimensions (measured from lateral radiographs). Multivariate statistical analysis of the skeletal variables in a group of 50 subjects led to the recognition of six independent factors determining facial shape, i.e., cranial base length, lower facial height, cranial base flexure and prognathism, facial width, mandibular length, and upper facial height. In 29 of these subjects, the cross-sectional areas of the jaw muscles were determined, and correlations between these areas and the scores on the above-mentioned factors were calculated. It appeared that the cross-sectional areas of temporalis and masseter muscles correlated positively with facial width, whereas the areas of masseter and both pterygoid muscles did so with mandibular length. It has been shown experimentally that a decrease in jaw muscle size in various animals likewise has an effect on facial width and mandibular length. Our results therefore support the hypothesis that in man too the jaw muscles affect facial growth and partly determine the final facial dimensions. They also hint that the role of each muscle is different.     

Variability of the Upper Palaeolithic skulls from Predmostí near Prerov (Czech Republic): craniometric comparison with recent human standards.

One of the largest skeletal series of the Upper Palaeolithic period from Predmostí was destroyed during the Second World War, but the study of this material continues up to the present. The discovery of Matiegka's original photographic documentation on glass plates [Velemínská et al., 2004. The use of recently re-discovered glass plate photo-documentation of those human fossil finds from Predmostí u Prerova destroyed during World War II. J. Nat. Mus. Nat. Hist. Ser. 173, 129-132] gives an opportunity to perform a new and detailed craniometric analysis of five adult skulls in their lateral projection. The craniometric data were analysed using specialised Craniometrics software, and the analysis included morphological and dimensional comparisons with current Central European norms. The aim of the study was not only to monitor the skull shape as a whole, but predominantly, to evaluate the size and shape of various parts of the splanchnocranium. The Upper Palaeolithic skulls are significantly longer, and male skulls are also higher than the current norms. The crania of anatomically modern humans are characterised by two general structural features: mid-lower facial retraction and neurocranial globularity. The height of the face of the Palaeolithic skulls corresponds to that of the current Central European population. The face has a markedly longer mandibular body (3-4 SD), while female mandibular rami are shorter. The skulls are further characterised by a smaller gonial angle, the increased steepness of the mandibular ramus, and the greater angle of the chin. These changes in the size and shape associated with anterior rotation of the face produce a strong protrusion of both jaws, but the sagittal inter-maxillary relationships remain unchanged. The observed facial morphology is similar to the Czech Upper Palaeolithic skulls from Dolní Vestonice. This study confirms the main diachronic changes between skulls of Upper Palaeolithic and present-day human populations.     

Linear mixed models for longitudinal shape data with applications to facial modeling

We present a novel application of methods for analysis of high-dimensional longitudinal data to a comparison of facial shape over time between babies with cleft lip and palate and similarly aged controls. A pairwise methodology is used that was introduced in Fieuws and Verbeke (2006) in order to apply a linear mixed-effects model to data of high dimensions, such as describe facial shape. The approach involves fitting bivariate linear mixed-effects models to all the pairwise combinations of responses, where the latter result from the individual coordinate positions, and aggregating the results across repeated parameter estimates (such as the random-effects variance for a particular coordinate). We describe one example using landmarks and another using facial curves from the cleft lip study, the latter using B-splines to provide an efficient parameterization. The results are presented in 2 dimensions, both in the profile and in the frontal views, with bivariate confidence intervals for the mean position of each landmark or curve, allowing objective assessment of significant differences in particular areas of the face between the 2 groups. Model comparison is performed using Wald and pseudolikelihood ratio tests.     

Polynesian face and dentition: functional perspective

Widely dispersed throughout the Pacific, Polynesians are a biologically distinctive people in form and size of both body and head. Large-bodied and well-muscled, their body phenotype is suited to life in a thermolabile oceanic environment. Their craniofacial skeleton is large and robust, with mandibular size and form (the "rocker" mandible) being especially characteristic. In this paper the Polynesian variants of body form, and of facial size (including dentition) and form, are interpreted from a functional perspective.     

Craniofacial morphology of Homo floresiensis: description, taxonomic affinities, and evolutionary implication

This paper describes in detail the external morphology of LB1/1, the nearly complete and only known cranium of Homo floresiensis. Comparisons were made with a large sample of early groups of the genus Homo to assess primitive, derived, and unique craniofacial traits of LB1 and discuss its evolution. Principal cranial shape differences between H. floresiensis and Homo sapiens are also explored metrically. The LB1 specimen exhibits a marked reductive trend in its facial skeleton, which is comparable to the H. sapiens condition and is probably associated with reduced masticatory stresses. However, LB1 is craniometrically different from H. sapiens showing an extremely small overall cranial size, and the combination of a primitive low and anteriorly narrow vault shape, a relatively prognathic face, a rounded oval foramen that is greatly separated anteriorly from the carotid canal/jugular foramen, and a unique, tall orbital shape. Whereas the neurocranium of LB1 is as small as that of some Homo habilis specimens, it exhibits laterally expanded parietals, a weak suprameatal crest, a moderately flexed occipital, a marked facial reduction, and many other derived features that characterize post-habilis Homo. Other craniofacial characteristics of LB1 include, for example, a relatively narrow frontal squama with flattened right and left sides, a marked frontal keel, posteriorly divergent temporal lines, a posteriorly flexed anteromedial corner of the mandibular fossa, a bulbous lateral end of the supraorbital torus, and a forward protruding maxillary body with a distinct infraorbital sulcus. LB1 is most similar to early Javanese Homo erectus from Sangiran and Trinil in these and other aspects. We conclude that the craniofacial morphology of LB1 is consistent with the hypothesis that H. floresiensis evolved from early Javanese H. erectus with dramatic island dwarfism. However, further field discoveries of early hominin skeletal remains from Flores and detailed analyses of the finds are needed to understand the evolutionary history of this endemic hominin species.     

Anthropometric analysis of the face in hypohidrotic ectodermal dysplasia: a family study

Sixteen individuals with hypohidrotic ectodermal dysplasia (HED) were compared to normal standards as well as to 16 unaffected family members by using a series of 20 anthropometric measurements of the head and face. Individuals with HED were generally smaller than normal controls or their unaffected relatives. However, this size reduction was not uniform. Instead, it was most evident in the anterior-posterior dimensions of the lower two-thirds of the face, in facial height, and in the size of the ears, nose, and mouth. A stepwise discriminant function analysis indicated that a function constructed from four variables (depth of the lower face, width of the nose, mandibular arc, and total facial height) could accurately classify 96.7% of the 32 individuals in the combined sample of affected and unaffected individuals. These findings demonstrated that the face of individuals with HED is unique and can be useful in its diagnosis. Additional studies are needed to determine if similar-though-less-pronounced facial abnormalities can be used to detect minimally affected gene carriers of this presumably X-linked condition.     

Protective buttressing of the hominin face

When humans fight hand‐to‐hand the face is usually the primary target and the bones that suffer the highest rates of fracture are the parts of the skull that exhibit the greatest increase in robusticity during the evolution of basal hominins. These bones are also the most sexually dimorphic parts of the skull in both australopiths and humans. In this review, we suggest that many of the facial features that characterize early hominins evolved to protect the face from injury during fighting with fists. Specifically, the trend towards a more orthognathic face; the bunodont form and expansion of the postcanine teeth; the increased robusticity of the orbit; the increased robusticity of the masticatory system, including the mandibular corpus and condyle, zygoma, and anterior pillars of the maxilla; and the enlarged jaw adductor musculature are traits that may represent protective buttressing of the face. If the protective buttressing hypothesis is correct, the primary differences in the face of robust versus gracile australopiths may be more a function of differences in mating system than differences in diet as is generally assumed. In this scenario, the evolution of reduced facial robusticity in Homo is associated with the evolution of reduced strength of the upper body and, therefore, with reduced striking power. The protective buttressing hypothesis provides a functional explanation for the puzzling observation that although humans do not fight by biting our species exhibits pronounced sexual dimorphism in the strength and power of the jaw and neck musculature. The protective buttressing hypothesis is also consistent with observations that modern humans can accurately assess a male's strength and fighting ability from facial shape and voice quality.     

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.     

A model of developmental canalization,applied to human cranial form

Developmental mechanisms that canalize or compensate perturbations of organismal development (targeted or compensatory growth) are widely considered a prerequisite of individual health and the evolution of complex life, but little is known about the nature of these mechanisms. It is even unclear if and how a “target trajectory” of individual development is encoded in the organism’s genetic-developmental system or, instead, emerges as an epiphenomenon. Here we develop a statistical model of developmental canalization based on an extended autoregressive model. We show that under certain assumptions the strength of canalization and the amount of canalized variance in a population can be estimated, or at least approximated, from longitudinal phenotypic measurements, even if the target trajectories are unobserved. We extend this model to multivariate measures and discuss reifications of the ensuing parameter matrix. We apply these approaches to longitudinal geometric morphometric data on human postnatal craniofacial size and shape as well as to the size of the frontal sinuses. Craniofacial size showed strong developmental canalization during the first 5 years of life, leading to a 50% reduction of cross-sectional size variance, followed by a continual increase in variance during puberty. Frontal sinus size, by contrast, did not show any signs of canalization. Total variance of craniofacial shape decreased slightly until about 5 years of age and increased thereafter. However, different features of craniofacial shape showed very different developmental dynamics. Whereas the relative dimensions of the nasopharynx showed strong canalization and a reduction of variance throughout postnatal development, facial orientation continually increased in variance. Some of the signals of canalization may owe to independent variation in developmental timing of cranial components, but our results indicate evolved, partly mechanically induced mechanisms of canalization that ensure properly sized upper airways and facial dimensions.     

Roentgencephalometric study of cranial interrelations

Lateral X-ray films of the skull obtained in 50 normal adult males were used for studies of correlations between 26 characteristics of the size, shape, and position of the face and nine characteristics of the neurocranium in all mutual combinations. The results disclosed that the relations between individual cranial components were regulated by certain principles. The correlations between size dimensions were mostly slight; a closer relationship showed some characteristics of the shape and position. The most important variable exerting an effect on the configuration of the skull as a whole represented the angle of the cranial base which produced the rotation of the neurocranium and the face and thus acted on a series of other correlations. Of some importance as well was the length of the mandibular ramus acting on the shape and position of the lower jaw and on the vertical maxillomandibular relations. The close relationship between the anteroposterior position of both jaws was due to compensation mechanisms rather than to the identical size of both jaws. On normalization of the disturbed saggital jaw relations, the dentoalveolar components of both jaws as well as the subalveolar component of the mandible participated equally. In vertical direction the lower face showed a certain developmental independence. The discussed interrelations formed the basis for studies of the mechanisms regulating the intracranial development and the changes occurring in various anomalies, as well as for understanding the compensation and adaptation abilities of individual cranial components.     

Hawaiian craniofacial morphometrics: Average Mokapuan skull,artificial cranial deformation,and the “rocker” mandible

Craniofacial morphology and cultural cranial deformation were analyzed by the computer morphometric system in 79 adult Hawaiian skulls from Mokapu, Oahu. The average Hawaiian male was large, but similar in shape to the female. Both were larger than the present Caucasian, showed a greater dental protrusion, and possessed a larger ANB angle, flatter cranial base, and larger facial heights. Correlations in Hawaiian craniofacial structure were found between an increasing mandibular plane angle and (1) shorter posterior facial height, (2) larger gonial angle, (3) larger cranial base angle, and (4) smaller SNA and SNB angles. Of the 79 skulls studied, 8. 9% were found to have severe head molding or intentional cranial deformation. Significant statistical differences between the molded group and the nonmolded group are, in decreasing significance: (1) larger upper face height, (2) smaller glabella to occiput distance, and (3) increased lower face height with deformation. The morphometric differences were readily seen by graphic comparison between groups. It is postulated that external forces to the neurocranium result in redirection of the growth vectors in the neurocranial functional matrix, including the cranial base, and secondarily, to the orofacial functional matrix. There is a possibility that the cranial deformation is a retention of the normal birth molding changes. The Polynesian “rocker jaw” was found in 81% to 95% of this populace. This mandibular form occurs only with attainment of adult stature and craniofacial form. This data agrees with the hypothesis that mandibular form is modified by the physical forces present and their direction in the orofacial functional matrix.     

Human occlusofacial form commonality: race-related traits, ontogeny, paedomorphism, and functional correlates

Previous cephalometric studies of occlusofacial angular and metric data show small variability within and between population samples of Caucasian children and adults. Comparable results were obtained by use of standardized X-ray cephalometry and direct craniometric measurements of skulls of mature specimens of Alouatta caraya and Cebus capucinus. These data measure facial and occlusofacial form and form relationships. Identical measuring techniques are applied to compare data derived from headplates of 19 young adult American Caucasians and 19 articulated Hindu skulls of mature individuals. The data are expanded to include measurements that relate symphyseal form to occlusal and nasomaxillary structure. The facial plane and an "internal" facial plane (nasion to a point tangential to the superior genial tubercles) straddle the mandibular symphysis. The angles they form at nasion and at their intersections with the maxillary occlusal plane were derived and statistically analyzed. The occlusal plane intersect angles show small absolute variability and are not significantly different (P greater than .05) in the two groups. The nasion plane angles are significantly different (P less than .025), reflecting sample differences in facial height. Linear measurements confirm a considerably larger dentoalveolar prognathism in the Hindu group. Data derived from the geometrically averaged tracings of Bjork, Broadbent, Sassouni, and Solow provide further evidence of occlusofacial form commonality. These shared phyletically conservative traits are evidence of highly integrated morphological adaptations for the attainment, maintenance, and functioning of the species-wide pattern of precisely fitting complementary crowns and crown parts that constitute the dental occlusion.     

Head proportion and shape of the head of children between 2 and 7 years--results of a longitudinal study

Based upon a longitudinal anthropometric study with integrated picture documentation, the head proportion and head shape of two to seven years old children will be analysed. The study was initiated in April 2002 in the administrative districts of Potsdam and Potsdam-Mittelmark (Germany). 351 boys and girls have been measured and photographed. The children shall be examined up to four years until September 2006 with a half year investigation distance. Twenty-three anthropometric dimensions have been measured including six dimensions of the head. These dimensions of the head are the head length, head breadth, auricular height, interpupillary distance, facial height and lower face height. Furthermore, digital pictures have been taken of each child. So the head of each child can be examined in a frontal view (Norma frontalis) and in a lateral view (Norma lateralis) and will be analysed relating to changing proportions of the head. The results demonstrated and discussed here are based on a longitudinal succession of photos and point out a method to make individual patterns of the changing head proportions and head shape visual on pictures.     

Strain in the galago facial skull

Little experimental work has been directed at understanding the distribution of stresses along the facial skull during routine masticatory behaviors. Such information is important for understanding the functional significance of the mammalian circumorbital region. In this study, bone strain was recorded along the dorsal interorbit, postorbital bar, and mandibular corpus in Otolemur garnettii and O. crassicaudatus (greater galagos) during molar chewing and biting. We determined principal-strain magnitudes and directions, compared peak shear-strain magnitudes between various regions of the face, and compared galago strain patterns with similar experimental data for anthropoids. This suite of analyses were used to test the facial torsion model (Greaves [1985] J Zool (Lond) 207:125-136; [1991] Zool J Linn Soc 101:121-129; [1995] Functional morphology in vertebrate paleontology. Cambridge: Cambridge University Press, p 99-115). A comparison of galago circumorbital and mandibular peak strains during powerful mastication indicates that circumorbital strains are very low in magnitude. This demonstrates that, as in anthropoids, the strepsirhine circumorbital region is highly overbuilt for countering routine masticatory loads. The fact that circumorbital peak-strain magnitudes are uniformly low in both primate suborders undermines any model that emphasizes the importance of masticatory stresses as a determinant of circumorbital form, function, and evolution. Preliminary data also suggest that the difference between mandibular and circumorbital strains is greater in larger-bodied primates. This pattern is interpreted to mean that sufficient cortical bone must exist in the circumorbital region to prevent structural failure due to nonmasticatory traumatic forces. During unilateral mastication, the direction of epsilon(1) at the galago dorsal interorbit indicates the presence of facial torsion combined with bending in the frontal plane. Postorbital bar principal-strain directions during mastication are oriented, on average, very close to 45 degrees relative to the skull's long axis, much as predicted by the facial torsion model. When chewing shifts from one side of the face to the other, there is a characteristic reversal or flip-flop in principal-strain directions for both the interorbit and postorbital bar. Although anthropoids also exhibit an interorbital reversal pattern, peak-strain directions for this clade are opposite those for galagos. The presence of such variation may be due to suborder differences in relative balancing-side jaw-muscle force recruitment. Most importantly, although the strain-direction data for the galago circumorbital region offer support for the occurrence of facial torsion, the low magnitude of these strains suggests that this loading pattern may not be an important determinant of circumorbital morphology.     

Bioanthropological analysis of human occipital condyles using geometric morphometric method

Sex differences are present in all parts of the body, including the skeletal system. Several methods are used to analyze the sex differences of skeleton, while more recently, a new method called geometric morphometry has been used. The aim of this study was to examine the sexual dimorphism of occipital condyles on human skulls originating from the population of Bosnia and Herzegovina using the geometric morphometric method.Material and methodsThe study was conducted on 214 human skulls of known gender from Bosnian population. For analysis of sexual dimorphism of occipital condyles, we used geometric morphometry, where all the skulls were scanned to obtain three-dimensional skull models. On the obtained models, we marked anthropometric points on occipital condyles in a Landmark Editor program from which we exported data in the form NTSYS file and analyzed it in MorphoJ program.ResultsFirst principal component PC1 describes 26.917% of total variability, the second principal component PC2 describes 20.992% of total variability, while the first eight principal components together describe 100% of total variability. The greatest variability between the male skulls and female skulls was present in the anterior-posterior diameter (length of occipital condyles). Discriminant functional analysis of the shape and size of the occipital condyles was possible with 69.50% accuracy for male skulls and with 60.27% accuracy for female skulls. The size of the occipital condyles showed a statistically significant effect on sexual determination. Discriminant functional analysis of the shape of the occipital condyles without affecting size enabled the determination of gender with with 65.96% accuracy for male skulls and with 63.01% accuracy for female skulls.ConclusionAnalysis of sexual dimorphism of occipital condyles using geometric morphometry showed statistically significant differences in the shape and size of occipital condyles between the sexes. The accuracy of sex determination based on occipital condyles was higher for male gender.