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PubMedDate not supplied

Developmental integration in a complex morphological structure: how distinct are the modules in the mouse mandible?

The mouse mandible has long served as a model system for studying the development and evolution of complex morphological structures. We used the methods of geometric morphometrics to reassess the hypothesis that the mandible consists of two separate modules: an anterior part bearing the teeth and a posterior part with muscle attachment surfaces and articulating with the skull. The analyses particularly focused on covariation of fluctuating asymmetry, because such covariation is due exclusively to direct interactions between the developmental processes that produce the traits of interest, whereas variation of traits among individuals also reflects other factors. The patterns of fluctuating asymmetry and individual variation were only partly consistent, indicating that developmental processes contribute differentially to variation at different levels. The results were in agreement with the hypothesis that the anterior and posterior parts of the mandible are separate develop-mental modules. Comparison of all alternative partitions of the landmarks into two contiguous subsets confirmed the hypothesis for the location of the boundary between modules but also underscored that the separation between them is not complete. Modularity is therefore manifest as the relative independence of parts within the framework of overall integration of the mandible as a whole-it is a matter of degrees, not all or nothing.

Analysis of Variance↗
PubMed2003

The shape of the Neandertal femur is primarily the consequence of a hyperpolar body form.

Neandertal femora are distinct from contemporaneous near-modern human femora. Traditionally, these contrasts in femoral shape have been explained as the result of the elevated activity levels and limited cultural abilities of Neandertals. More recently, however, researchers have realized that many of these femoral differences may be explained by the cold-adapted bodies of Neandertals vs. the warm-adapted bodies of near-modern humans. This study explicitly tests this proposed link between climate-induced body proportions and femoral shape by considering the entire hip as a unit by using geometric morphometric methods adapted to deal with articulated structures. Based on recent human patterns of variation, most contrasts in shape between the femora of Neandertals and near-modern humans seem to be secondary consequences of differences in climate-induced body proportions. These results, considered in light of hip mechanics during growth, highlight the importance of developmental and functional integration in determining skeletal form.

Animals↗
PubMed2003

Encephalization and allometric trajectories in the genus Homo: evidence from the Neandertal and modern lineages.

The term "encephalization" is commonly used to describe an enlargement in brain size, considered as either absolute endocranial volumes or relative values in relation to body size. It is widely recognized that a considerable endocranial expansion occurred throughout the evolution of the genus Homo. This article aims to evaluate whether this phenomenon was the outcome of distinct evolutionary lineages, reaching similar brain expansions but through different trajectories. Endocranial morphology was studied in a sample of fossil hominines by multivariate approaches using both traditional metrics and geometric morphometrics. The analysis was focused on the transition from a generalized archaic pattern within the genus Homo to the modern morphology and compared with changes that occurred along the Neandertal lineage. The main result was the identification of two different evolutionary trajectories, in which a similar expansion in endocranial size has been reached by different changes in shape. Along the Neandertal lineage we observed maintenance of an "archaic" endocranial model, in which a large amount of variability is based on a single allometric trend. By contrast, when modern endocasts were compared with nonmodern ones, we found important differences apparently led by a parietal expansion. In this light, the origin of our species may have represented the opportunity to surpass the constraints imposed on encephalization by the ontogenetic pattern shared by nonmodern Homo representatives.

Animals↗
PubMed2000

Use of two-block partial least-squares to study covariation in shape.

The relatively new two-block partial least-squares method for analyzing the covariance between two sets of variables is described and contrasted with the well-known method of canonical correlation analysis. Their statistical properties, type of answers, and visualization techniques are discussed. Examples are given to show its usefulness in comparing two sets of variables--especially when one or both of the sets of variables are shape variables from a geometric morphometric study.

Analysis of Variance↗
PubMed2003

The geometry of the marmot (rodentia: sciuridae) mandible: phylogeny and patterns of morphological evolution.

Marmots have a prominent role in the study of mammalian social evolution, but only recently has their systematics received the attention it deserves if sociobiological studies are to be placed in a phylogenetic context. Sciurid morphology can be used as model to test the congruence between morphological change and phylogeny because sciurid skeletal characters are considered to be inclined to convergence. However, no morphological study involving all marmot species has ever been undertaken. Geometric morphometric techniques were applied in a comparative study of the marmot mandible. The adults of all 14 living marmot species were compared, and mean mandible shape were used to investigate morphological evolution in the genus Marmota. Three major trends were observed. First, the phylogenetic signal in the variation of landmark geometry, which describes mandible morphology, seems to account for the shape differences at intermediate taxonomic levels. The subgenera Marmota and Petromarmota, recently proposed on the basis of mitochondrial cytochrome b sequence, receive support from mandible morphology. When other sciurid genera were included in the analysis, the monophyly of the genus Marmota and that of the tribe Marmotini (i.e., marmots, prairie dogs, and ground squirrels) was strengthened by the morphological data. Second, the marmotine mandible may have evolved as a mosaic of characters and does not show convergence determined by size similarities. Third, allopatric speciation in peripheral isolates may have acted as a powerful force for modeling shape. This hypothesis is strongly supported by the peculiar mandible of M. vancouverensis and, to a lesser degree, by that of M. olympus, both thought to have originated as isolated populations in Pleistocene ice-free refugia.

Adaptation, Biological↗
PubMed2003

The effect of phylogeny on interspecific body shape variation in darters (Pisces: Percidae).

We conducted a geometric morphometric analysis of interspecific body shape variation among representatives of 31 species of darters (Pisces: Percidae) to determine whether there is evidence of a phylogenetic effect in body shape variation. Cartesian transformation grids representing relative shape differences of individual species and subspecies revealed qualitative similarities within most traditionally recognized taxonomic groups (genera and subgenera). Canonical variates analysis and a UPGMA cluster analysis were conducted to explore further the relationships among body shapes of species; both analyses revealed patterns of variation consistent with the interpretation that shape is associated with taxonomic affinities. Normalized Mantel statistics revealed a significant positive association between body shape differences and phylogenetic interrelatedness for each of four recent phylogenetic hypotheses, providing evidence of a phylogenetic effect. This result is somewhat surprising, however, given the largely incompatible nature of these four phylogenies. We provide evidence that this result may be due to (1) the inclusion of multiple sets of closely related species to represent the traditionally recognized genera and subgenera within each phylogeny and/or (2) the inclusion of several species with relatively divergent shapes and their particular positions within the phylogenies relative to one another or to the other species of darters.

Animals↗
PubMed2006

Natural selection on Erysimum mediohispanicum flower shape: insights into the evolution of zygomorphy.

Paleontological and phylogenetic studies have shown that floral zygomorphy (bilateral symmetry) has evolved independently in several plant groups from actinomorphic (radially symmetric) ancestors as a consequence of strong selection exerted by specialized pollinators. Most studies focused on unraveling the developmental genetics of flower symmetry, but little is known about the adaptive significance of intraspecific flower shape variation under natural conditions. We provide the first evidence for natural selection favoring zygomorphy in a wild population of Erysimum mediohispanicum (Brassicaceae), a plant showing extensive continuous variation in flower shape, ranging from actinomorphic to zygomorphic flowers. By using geometric morphometric tools to describe flower shape, we demonstrate that plants bearing zygomorphic flowers received more pollinator visits and had the highest fitness, measured not only by the number of seeds produced per plant but also by the number of seeds surviving to the juvenile stage. This study provides strong evidence for the existence of significant fitness differences associated with floral shape variation in E. mediohispanicum, thus illuminating a pathway for the evolution of zygomorphy in natural populations.

Animals↗
PubMed2001

Facial growth: separating shape from size.

Optical surface scanning technologies produce dense three-dimensional (3D) data sets, which allow detailed analysis of surface morphology. This paper describes a method of analysing change in facial shape independently of change in size. The 3D data from three male subjects from the age of 6-21 years were recorded using an optical surface scanner. A series of 22 conventional landmarks were located with the aid of horizontal and vertical profiles across the face, and were analysed using geometric morphometrics. The 3D landmark co-ordinates were scaled and aligned using Generalized Procrustes Analysis (GPA) and analysed by Principal Component Analysis (PCA) to determine the shape change over the growth period for each individual. The results show that the centroid size reaches a steady value at different times for each of the subjects. When analysing shape versus age, highly significant correlations were found with principal component 1 (PC-1), but not with other principal components. PC-1 encompassed 40 per cent of the total variance for each subject. The movement of facial landmarks with time that is represented by PC-1 in each of the individuals is described. The use of these techniques has enabled the individual characteristics of facial growth to be identified and also has revealed the subtle changes in shape that continue after change in size has ceased.

Adolescent↗
PubMed2003

Size and shape measurement in contemporary cephalometrics.

The traditional method of analysing cephalograms--conventional cephalometric analysis (CCA)--involves the calculation of linear distance measurements, angular measurements, area measurements, and ratios. Because shape information cannot be determined from these 'size-based' measurements, an increasing number of studies employ geometric morphometric tools in the cephalometric analysis of craniofacial morphology. Most of the discussions surrounding the appropriateness of CCA, Procrustes superimposition, Euclidean distance matrix analysis (EDMA), thin-plate spline analysis (TPS), finite element morphometry (FEM), elliptical Fourier functions (EFF), and medial axis analysis (MAA) have centred upon mathematical and statistical arguments. Surprisingly, little information is available to assist the orthodontist in the clinical relevance of each technique. This article evaluates the advantages and limitations of the above methods currently used to analyse the craniofacial morphology on cephalograms and investigates their clinical relevance and possible applications.

Algorithms↗
PubMed2005

Maxillofacial growth in children with unilateral cleft lip and palate following secondary alveolar bone grafting: an interim evaluation.

This study evaluates the effect of alveolar bone grafting on the maxillofacial growth in children of mixed dentition with unilateral complete cleft lip-cleft palate. Twenty patients received an iliac crest alveolar bone graft between the ages of 6 years 10 months and 10 years 10 months, whereas 20 matched controls between the ages of 6 years 11 months and 10 years 6 months did not. Geometric morphometric assessments were used to localize alterations between the initial and final cephalographs in the two groups of cleft children, using Procrustes analysis and thin-plate spline analysis, in addition to conventional cephalometric techniques. It is concluded that no statistically significant difference in maxillofacial growth was found between the cleft children having received secondary alveolar bone grafting and the nongraft controls in general during the first to third postoperative years. Further investigation will be undertaken to determine the long-term effects after the age of skeletal maturity.

Alveolar Bone Loss↗
PubMed1998

Left-right asymmetry of fly wings and the evolution of body axes.

The body plan of Drosophila, and presumably that of other insects, develops under the control of anterio-posterior and dorsal ventral axes, but no evidence for a left-right axis has yet been found. We used geometric morphometrics to study the wings in three species of flies: Drosophila melanogaster, Musca domestica and Glossina palpalis gambiensis. In all three species, we found that both size and shape showed subtle, but statistically significant directional asymmetry. For size, these asymmetries were somewhat inconsistent within and between species, but for shape, highly significant directional asymmetry was found in all samples examined. These systematic left-right differences imply the existence of a left-right axis that conveys distinct positional identities to the wing imaginal discs on either body side. Hence, the wing discs of Drosophila may be a new model to study the developmental genetics of left-right asymmetry. The asymmetries of shape were similar among species, suggesting that directional asymmetry has been evolutionarily conserved since the three lineages diverged. We discuss the implications of this evolutionary conservatism in conjunction with results from earlier studies that showed a lack of genetic variation for directional asymmetry in Drosophila.

Animals↗
PubMed2000

Independence between developmental stability and canalization in the skull of the house mouse.

The relationship between the two components of developmental homeostasis, that is canalization and developmental stability (DS), is currently debated. To appraise this relationship, the levels and morphological patterns of interindividual variation and fluctuating asymmetry were assessed using a geometric morphometric approach applied to the skulls of laboratory samples of the house mouse. These three samples correspond to two random-bred strains of the two European subspecies of the house mouse and their F1 hybrids. The inter- and intraindividual variation levels were found to be smaller in the hybrid group compared to the parental ones, suggesting a common heterotic effect on skull canalization and DS. Both buffering mechanisms might then depend on the same genetic condition, i.e. the level of heterozygosity. However, related morphological patterns did not exhibit any congruence. In contradiction with previous studies on insect wing traits, we therefore suggest that canalization and DS may not act on the same morphological characters. The fact that this discrepancy could be related to the functional importance of the symmetry of the characters under consideration is discussed in the light of our knowledge of the genetic bases of both components of developmental homeostasis.

Animals↗
PubMed2002

Visualizing patterns of craniofacial shape variation in Homo sapiens.

The geometric morphometric analysis of shape variation in complex biological structures such as the human skull poses a number of specific challenges: the registration of homologous morphologies, the treatment of bilateral symmetry, the graphical representation of form variability in three dimensions and the interpretation of the results in terms of differential growth processes. To visualize complex patterns of shape change, we propose an alternative to classical Cartesian deformation grids in the style of D'Arcy W. Thompson. Reference to the surface structures of the organism under investigation permits a comprehensive visual grasp of shape change and its tentative interpretation in terms of differential growth. The application of this method to the analysis of human craniofacial shape variation reveals distinct modes of growth and development of the neurocranial and viscerocranial regions of the skull. Our data further indicate that variations in the orientation of the viscerocranium relative to the neurocranium impinge on the shapes of the face and the cranial vault.

Adolescent↗
PubMed2006

Developmental response to cold stress in cranial morphology of Rattus: implications for the interpretation of climatic adaptation in fossil hominins.

Adaptation to climate occupies a central position in biological anthropology. The demonstrable relationship between temperature and morphology in extant primates (including humans) forms the basis of the interpretation of the Pleistocene hominin Homo neanderthalensis as a cold-adapted species. There are contradictory signals, however, in the pattern of primate craniofacial changes associated with climatic conditions. To determine the direction and extent of craniofacial change associated with temperature, and to understand the proximate mechanisms underlying cold adaptations in vertebrates in general, dry crania from previous experiments on cold- and warm-reared rats were investigated using computed tomography scanning and three-dimensional digitization of cranial landmarks. Aspects of internal and external cranial morphology were compared using standard statistical and geometric morphometric techniques. The results suggest that the developmental response to cold stress produces subtle but significant changes in facial shape, and a relative decrease in the volume of the maxillary sinuses (and nasal cavity), both of which are independent of the size of the skull or postcranium. These changes are consistent with comparative studies of temperate climate primates, but contradict previous interpretations of cranial morphology of Pleistocene Hominini.

Adaptation, Biological↗
PubMed2000

Morphological intergration between development compartments in the Drosophila wing.

Developmental integration is the covariation among morphological structures due to connections between the developmental processes that built them. Here we use the methods of geometric morphometrics to study integration in the wing of Drosophila melanogaster. In particular, we focus on the hypothesis that the anterior and posterior wing compartments are separate developmental units that vary independently. We measured both variation among genetically diverse individuals and random differences between body sides of single individuals (fluctuating asymmetry, FA). For both of these sources of variation, the patterns of variation identified by principal component analyses all involved landmarks in both the anterior and posterior compartments simultaneously. Analyses focusing exclusively on the covariation between the anterior and posterior compartments, by the partial least-squares method, revealed pervasive integration of the two compartments, for both individual variation and FA. These analyses clearly indicate that the anterior and posterior compartments are not separate units of variation, but that the covariation between compartments is sufficient to account for nearly all the variation throughout the entire wing. We conclude that variation among individuals as well as the developmental perturbations responsible for FA generate shape variation primarily through developmental processes that are integrated across both compartments. In contrast, much less of the shape variation in our sample can be attributed to the localized processes that establish the identity of particular wing veins.

Animals↗
PubMed2001

Quantitative genetics of geometric shape in the mouse mandible.

We combine the methods of geometric morphometrics and multivariate quantitative genetics to study the patterns of phenotypic and genetic variation of mandible shape in random-bred mice. The data are the positions of 11 landmarks on the mandibles of 1,241 mice from a parent-offspring breeding design. We use Procrustes superimposition to extract shape variation and restricted maximum likelihood to estimate the additive genetic and environmental components of variance and covariance. Matrix permutation tests showed that the genetic and phenotypic as well as the genetic and environmental covariance matrices were similar, but not identical. Likewise, principal component analyses revealed correspondence in the patterns of phenotypic and genetic variation. Patterns revealed in these analyses also showed similarities to features previously found in the effects of quantitative trait loci and in the phenotypes generated in gene knockout experiments. We used the multivariate version of the breeders' equation to explore the potential for short-term response to selection on shape. In general, the correlated response is substantial and regularly exceeds the direct response: Selection applied locally to one landmark usually produces a response in other parts of the mandible as well. Moreover, even selection for shifts of the same landmark in different directions can yield dramatically different responses. These results demonstrate the role of the geometry and anatomical structure of the mandible, which are key determinants of the patterns of the genetic and phenotypic covariance matrices, in molding the potential for adaptive evolution.

Animals↗
PubMed2003

Allometric and nonallometric components of Drosophila wing shape respond differently to developmental temperature.

Phenotypic plasticity of wing size and shape of Drosophila simulans was analyzed across the entire range of viable developmental temperatures with Procrustes geometric morphometric method. In agreement with previous studies, size clearly decreases when temperature increases. Wing shape variation was decomposed into its allometric (24%) and nonallometric (76%) components, and both were shown to involve landmarks located throughout the entire wing blade. The allometric component basically revealed a progressive, monotonous variation along the temperature. Surprisingly, nonallometric shape changes were highly similar at both extremes of the thermal range, suggesting that stress, rather than temperature per se, is the key developmental factor affecting wing shape.

Analysis of Variance↗
PubMed2005

Midfacial and mandibular morphometry of children with Class II and Class III malocclusions.

There is insufficient evidence in conventional cephalometric analysis of the actual sites of putative maxillofacial change in Class II and Class III malocclusions. The purpose of this study was to provide more information about the morphological characteristics of the midfacial complex and mandible in children with Class II or III malocclusions. Seventy children with Class II, division 1 malocclusion and 70 children with Class III malocclusion were compared with 70 children with normal occlusion. This study was conducted to carry out geometric morphometric assessments to localize alterations using Procrustes analysis and thin-plate spline analysis. Procrustes analysis indicated the midfacial and mandibular morphologies differed between normal occlusion subjects and subjects with Class II or Class III malocclusion (P<0.0001). The deformations in subjects with Class II malocclusion may represent a developmental elongation of the palatomaxillary complex and a shortening of the mandible anteroposteriorly, which leads to the appearance of a protruding midface and retruding mandibular profile. In contrast, the deformations in subjects with Class III malocclusion may represent a developmental shortening of the palatomaxillary complex and elongation of the mandible anteroposteriorly, which leads to the appearance of a retrognathic midface and prognathic mandibular profile.

Cephalometry↗