Craniofacial growth of fetal Macaca nemestrina: A cephalometric roentgenographic study

The prenatal growth of the macaque craniofacial skeleton is described using lateral radiographs of 82 fetal and 25 neonatal Macaca nemestrina whose known gestational ages range from 50 to 186 days. The ossification sequence of the craniofacial bones resembles that in the human fetus. During gestation, the macaque neurocranium loses its round, globular shape, becoming flattened and elongated in an anteroposterior direction. In contrast, the morphologic pattern of the face is established early in fetal life, and little change takes place during the remaining prenatal period. The macaque craniofacial dimensions develop along the general skeletal growth pattern, unlike the human craniofacial dimensions, which follow an intermediate pattern between the neural and general skeletal patterns. However, despite minor differences, the macaque and human fetal faces follow the same basic patterns of growth.     

A method of deriving subgroups of a population: a study of craniofacial taxonomy

Subgroups within a population are often difficult to discover and describe except by subjective methods. In this study, cluster analysis (numerical taxonomy) methods were used on selected craniofacial measurements obtained from 308 North American White children of both sexes in the age range 6–18 to derive categories of skeletal facial types. Two different cluster analysis approaches were used in conjunction with a separate overall evaluation of facial balance, an independent measure of maxillo-mandibular relationship (AB/FOP), and a traditional classification (Angle). The categories derived rest on corroborative and overlapping evidence from each of those methods. The categories were examined to determine if it is possible to classify a sample by means of cluster analysis, the size and discreteness of each class, how they compare with the Angle classification of the same sample, and the percentage of individuals that may be identified unequivocally by this classification scheme. Five categories were obtained. Labeled Category A — Category E, they show some correspondence to the Angle Classes I, II, and III, but categories A, B, and C appear to be subgroups, heretofore undetected, of Angle Class II. Categories D and E correspond to Angle Classes I and III, respectively. The categories are more realistic and informative than the Angle classes. Each category is reported along with its distinguishing skeletal characteristics.     

A brief note on growth physiology of plants

The concepts of physiological structure of plant axial organ and its main components are discussed. Their physiological meanings, in particular the role of the surface and the xylem proton pumps are highlighted: the former loosens the cell wall via acidification, and the latter produces the driving force for active uptake of water. Theoretical and experimental examination on the validity of the Lock-hart growth equation is reviewed. Development of a new experimental system, perfusible glycerinated hollow cylinder of cowpea hypocotyl, demonstrates the validity of the Lock-hart type mechanical equation even in such anin vitro system. The pH-dependency of both extensibility and yield threshold offer a strong support for the acid growth theory. A molecular model of cell wall extension is proposed on the basis of these results. The importance of growth regulation via control of the cell wall yield threshold is demonstrated as a very economical way, by an analogy with the performance of electron tube of triode type. Also augmentation of the classic acid growth theory is proposed on the basis of Katou's diagram and the Katou-Furumoto's model of active water uptake.     

Comparative craniofacial variation in Navajo Indians and North American Caucasians

Landmarks digitized from lateral cephalometric radiographs of 107 Navajo Indians between 10 and 12 years of age were analyzed to determine coefficients of variation or standard deviations for 38 cephalometric measurements. These values were compared with the same measures of variation for identical measurements on North American whites derived from the Michigan and Philadelphia Growth Studies. For the majority of variables, there were no differences between groups. Variation for the genetically and environmentally isolated Navajo Indians was the same as that of the highly diverse Caucasian samples. However, measurements of upper, lower, and total anterior facial height (N-ANS, ANS-Me, and N-Me, respectively) indicate that these features are substantially less variable in Navajo Indians relative to the Michigan and Philadelphia populations.     

Age factor and the pattern of change in craniofacial structures

Today special emphasis is being placed upon the understanding of human aging and this study is an attempt to shed light on the craniofacial complex during the later years. Postcranial skeletal alteration is clear and it is now evident that cranial and facial structures are no exception to the aging process. The longitudinal information presented here indicates continuing overall growth from early adulthood to later life. The cranium thickens and the skull diameter increases. Endocranial dimensions enlarge as well. This suggests larger overall skull size and expansion of the cranial cavity. The visceral cranial structures also participate in the continuing growth process. Enlargement in all areas seems to be of similar magnitude except for skull thickness, sella turcica, and frontal sinus. The size increase in these three structures is greater than in other segments examined. In essence, the craniofacial complex is in a state of growth throughout life. The entire system is involved in a process of symmetrical enlargement.     

Multivariate analysis of craniofacial measurements in twin and family data

Thirty-three cephalometric variables and height have been measured on each of 630 individuals (316 male and 314 female) from 157 families. After age and sex differences were adjusted for each measurement, a rotated factor analysis was undertaken to account for the variation by a limited number of linear combinations of the adjusted measurements. It was found that most of the variation could be explained by nine independent factors. Finally, correlation coefficients were computed on twins, siblings and parent-offspring data for factor scores. The results suggest that each factor which is measured by a linear combination of a set of variables could result from the interaction of independent sets of genes with the environment.     

Characterization of craniofacial sutures using the finite element method

Characterizing the biomechanical behavior of sutures in the human craniofacial skeleton (CFS) is essential to understand the global impact of these articulations on load transmission, but is challenging due to the complexity of their interdigitated morphology, the multidirectional loading they are exposed to and the lack of well-defined suture material properties. This study aimed to quantify the impact of morphological features, direction of loading and suture material properties on the mechanical behavior of sutures and surrounding bone in the CFS. Thirty-six idealized finite element (FE) models were developed. One additional specimen-specific FE model was developed based on the morphology obtained from a µCT scan to represent the morphological complexity inherent in CFS sutures. Outcome variables of strain energy (SE) and von Mises stress (σ_vm) were evaluated to characterize the sutures’ biomechanical behavior. Loading direction was found to impact the relationship between SE and interdigitation index and yielded varied patterns of σ_vm in both the suture and surrounding bone. Adding bone connectivity reduced suture strain energy and altered the σ_vm distribution. Incorporating transversely isotropic material properties was found to reduce SE, but had little impact on stress patterns. High-resolution µCT scanning of the suture revealed a complex morphology with areas of high and low interdigitations. The specimen specific suture model results were reflective of SE absorption and σ_vm distribution patterns consistent with the simplified FE results. Suture mechanical behavior is impacted by morphologic factors (interdigitation and connectivity), which may be optimized for regional loading within the CFS.     

Keratan sulfate expression during avian craniofacial morphogenesis

Summary Using the monoclonal antibody MZ15 in immunocytochemical and ultrastructural studies we have been able to determine the spatiotemporal pattern of keratan sulfate (KS) distribution during quail craniofacial morphogenesis. KS-containing proteoglycans are found associated with invaginating placodes (olfactory, lens and otic), in developing pronephric tubules, notochord, pharynx and endocardium, and display developmental regulation. The appearance of such proteoglycans (PGs) during placode morphogenesis is particularly striking and we suggest that they may be an important component of the extracellular matrix which has been previously implicated in mediating the morphogenetic interactions and cell movements occurring at these sites. The otic vesicle during stage 18–22 displays a notable asymmetric distribution of KS-containing PGs. The role that these molecules may play and the reasons for this regionalization are, as yet, unclear but it is conceivable that the distribution of proteoglycans at this stage reflects subsequent differentiative events during otocyst development. Furthermore, our ultrastructural observations indicate that over the developmental period studied (H & H stages 8–22) keratan sulfate exists in at least two proteoglycan forms. Some spatiotemporal correlation has been found to exist between the distributions of KS-containing PGs and type II collagen as previously reported by Thorogood et al. (1986). We suggest that the proteoglycan detected at such sites is cartilage-specific proteoglycan and that it plays an important role, together with type II collagen, in the “signalling” mechanism which specifies the subsequent pattern of the chondrocranium. It is proposed that this interaction at epithelio-mesenchymal interfaces in the developing head parallels the matrix-mediated tissue interaction between notochord and somites which results in the formation of the cartilaginous primordia of the vertebrae from the sclerotomes as reported by Lash and Vasan (1978).     

Age differences in craniofacial dimensions among adults from Indian Knoll,Kentucky

Adult craniofacial expansion with aging has recently been documented in a living US white population sample (Israel, '73a, '77). The present study extends these findings to a prehistoric Amerindian skeletal sample from the Indian Knoll, Kentucky site. Sixteen craniofacial dimensions available for 136 adult males were compared in younger (20–34 years) and older (35–50 years) age groups. Of these, six dimensions showed a significant difference between age groups; all significantly different dimensions were larger in the older adult age group. The multivariate (T²) difference between age groups was highly significant. Comparison of results before and after a size standardization indicated that the majority of differences between age groups were associated with an overall size increase, or expansion with aging, and did not represent merely remodeling, or “shape” changes. The pattern of craniofacial change with age appeared generally similar to that observed in the modern US white sample; however, some differences were noted. It is shown that the age trends observed at Indian Knoll are most likely to reflect true craniofacial growth in size among the adult male inhabitants of the site, rather than secular trends or other artifacts of the sampling procedure. The causes for continuing adult craniofacial expansion are unknown, and probably involve a complex interaction of many factors. However, this pattern of change with age among adults does appear to be characteristic of population samples of widely differing genetic and environmental backgrounds.     

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.     

New directions in craniofacial morphogenesis

The vertebrate head is an extremely complicated structure: development of the head requires tissue-tissue interactions between derivates of all the germ layers and coordinated morphogenetic movements in three dimensions. In this review, we highlight a number of recent embryological studies, using chicken, frog, zebrafish and mouse, which have identified crucial signaling centers in the embryonic face. These studies demonstrate how small variations in growth factor signaling can lead to a diversity of phenotypic outcomes. We also discuss novel genetic studies, in human, mouse and zebrafish, which describe cell biological mechanisms fundamental to the growth and morphogenesis of the craniofacial skeleton. Together, these findings underscore the complex interactions leading to species-specific morphology. These and future studies will improve our understanding of the genetic and environmental influences underlying human craniofacial anomalies.     

Reflections on the face of Japan: a multivariate craniofacial and odontometric perspective

Craniofacial variables for modern and prehistoric Japanese were subjected to multivariate analysis to test the relationships of the people of Japan with mainland Asian and Oceanic samples. The modern Japanese are tied to Koreans, Chinese, Southeast Asians, and the Yayoi rice agriculturalists who entered Japan in 300 B.C. Together they make up a Mainland-Asia cluster of related populations. The prehistoric Jomon foragers, the original inhabitants of the Japanese archipelago, are the direct ancestors of the modern Ainu, who made a recognizable contribution to the warrior class--the Samurai--of feudal Japan. Together, they are associated with Polynesians and Micronesians in a Jomon-Pacific cluster of related populations. Jomon-to-Ainu tooth size reduction proceeded at the same rate as that observable in the post-Pleistocene elsewhere in the Old World.     

Lysyl oxidase-like 3b is critical for cartilage maturation during zebrafish craniofacial development

Vertebrate craniofacial development requires coordinated morphogenetic interactions between the extracellular matrix (ECM) and the differentiating chondrocytes essential for cartilage formation. Recent studies reveal a critical role for specific lysyl oxidases in ECM integrity required for embryonic development. We now demonstrate that loxl3b is abundantly expressed within the head mesenchyme of the zebrafish and is critically important for maturation of neural crest derived cartilage elements. Histological and ultrastructural analyses of cartilage elements in loxl3b morphant embryos reveal abnormal maturation of cartilage and altered chondrocyte morphology. Spatiotemporal analysis of craniofacial markers in loxl3b morphant embryos shows that cranial neural crest cells migrate normally into the developing pharyngeal arches but that differentiation and condensation markers are aberrantly expressed. We further show that the loxl3b morphant phenotype is not due to P53 mediated cell death but likely to be due to reduced chondrogenic progenitor cell proliferation within the pharyngeal arches. Taken together, these data demonstrate a novel role for loxl3b in the maturation of craniofacial cartilage and can provide new insight into the specific genetic factors important in the pathogenesis of craniofacial birth defects.     

The effects of hybridization on growth allometry and craniofacial form in Sulawesi macaques

The present research investigates the effects of hybridization between Macaca maurus and M. tonkeana on adult male form and patterns of growth allometry. Comparisons of adult hybrid mean phenotypic values with the adult averages of the parental species indicate a condition of heterosis for cranial vault length and crown-rump length. Negative heterosis is indicated for body mass. Regression parameters describing growth allometry are generated for four craniofacial measurement variables and one body measurement using both least squares and reduced major axis regression. Comparisons of hybrid and parental regression slopes and intercepts using analysis of covariance and t-tests suggest that there is a hybrid pattern of growth allometry characterized by an increase in regression slope values coupled with lower intercept values compared to those of the parental species and the parental averages for most regression parameters. Multivariate analyses of the adult and ontogenetic morphometric data indicate significant differences across species taxa in form and shape during development and adulthood. Our finding of significant differences between hybrids and their parental taxa in growth allometry and craniofacial form and shape during development challenges the assumption often made regarding the reproductive and taxonomic significance of observed ontogenetic divergence between Neandertals and modern humans. We propose that anthropological primatology, with its goal of developing nonhuman primate models for investigating human evolution, can provide a biologically relevant means by which to empirically estimate the taxonomic significance of morphological and ontogenetic divergence observed in the hominid fossil record.     

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.     

A note on the effects of co-mingling piglet litters on pre-weaning growth, injuries and responses to behavioural tests

The purpose of this study was to determine how co-mingling litters affected piglets’ pre-weaning growth, ear injuries, suckling behaviour and responses to behavioural tests used to measure coping abilities. Thirty sows and their respective litters were housed in standard farrowing crates until day 13 after birth. On day13, the partition between two neighbouring pens was removed for 20 litters allowing piglets to interact (forming 10 co-mingled litters). The remaining 10 control litters were kept in standard farrowing crates throughout the experiment. Three focal piglets from each litter were used for data collection. Focal piglets were weighed and ear injuries recorded on days 2, 4, 9, 12, 15 and 18 after birth. There were no differences in piglets’ weight gain before or after co-mingling. Ear injuries were more abundant in co-mingled litters on day 15 (P < 0.05) but these differences disappeared by day 18. Suckling behaviour was recorded on days 5, 8, 10, 14, 16 and 18 after birth. There were no differences in teat fidelity, suckling frequency and mother fidelity between treatments. Three behavioural tests, social challenge, isolation, and backtest, were performed before and after co-mingling. There were no treatment effects on piglets’ response to the isolation test and backtest. Co-mingled piglets showed longer latency for the first aggressive interaction (P < 0.05), spent more time in proximity to one another (P < 0.05) and performed less single bites (P < 0.05) than control piglets during the social challenge. In addition, the duration and frequency of aggressive interactions (P < 0.05) were lower in co-mingled piglets than control piglets. Co-mingling did not affect the frequency of single head thrusts or oral–nasal contact, but did tend to increase the frequency of escape attempts (P < 0.10). Our results suggest that co-mingling litters during lactation affects piglets’ social behaviour, by primarily decreasing aggressive interactions during social challenges.