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PubMed2003

A life-course perspective on social exclusion and poverty.

This article assesses whether it is possible to reconceptualize the traditional research approaches to the relationship between poverty and the life cycle on the basis of different sociological perspectives on the life course found in the literature. While the family-cycle approach, which was originally formulated by Seebohm Rowntree (1902), is criticized for being static, descriptive, normative and inflexible, dynamic poverty research is mostly confined to the quantitative analysis of income trajectories, and thus offers only a partial solution to our problem. However, the life-course perspective allows us to combine the best elements of these traditional approaches and to reconceptualize them into a general framework for the study of social exclusion and poverty. To this end, three sociological perspectives on the life course are considered: the traditional North-American life-course perspective formulated by Elder (1974), the Continental institutional approach, and a combined approach which we label the 'political economy of the life course'. Drawing from these three perspectives, we propose a general framework of analysis and formulate hypotheses regarding the phenomena of social exclusion and poverty over the life course which can subsequently be empirically validated.

Cultural Deprivation↗

PubMed2002

Relationship of lipid rafts to transient confinement zones detected by single particle tracking.

We examined the physical and chemical characteristics of transient confinement zones (TCZs) that are detected in single particle trajectories of molecules moving within the membrane of C3H 10T1/2 murine fibroblasts and their relationship to "rafts." We studied the lateral movement of different membrane molecules thought to partition to varying degrees into or out of the putative lipid domains known as rafts. We found that lipid analogs spend significantly less time in TCZs compared with Thy-1, a glycosylphosphatidylinositol-anchored protein, and GM1, a glycosphingolipid. For Thy-1, we found that zone abundance was markedly reduced by cholesterol extraction, suggesting that a major source of the observed temporary confinement is related to the presence of raft domains. More detailed analysis of particle trajectories reveals that zones can be revisited even tens of seconds after the original escape and that diffusion within the zones is reduced by a factor of approximately 2, consistent with the zone being a cholesterol-rich liquid-ordered phase. Surprisingly, transient confinement was not strongly temperature dependent. Overall, our data demonstrate that there are raft-related domains present in certain regions of the plasma membrane of C3H cells, which can persist for tens of seconds.

Animals↗

PubMed2006

Multiscale modeling of nucleosome dynamics.

Nucleosomes form the fundamental building blocks of chromatin. Subtle modifications of the constituent histone tails mediate chromatin stability and regulate gene expression. For this reason, it is important to understand structural dynamics of nucleosomes at atomic levels. We report a novel multiscale model of the fundamental chromatin unit, a nucleosome, using a simplified model for rapid discrete molecular dynamics simulations and an all-atom model for detailed structural investigation. Using a simplified structural model, we perform equilibrium simulations of a single nucleosome at various temperatures. We further reconstruct all-atom nucleosome structures from simulation trajectories. We find that histone tails bind to nucleosomal DNA via strong salt-bridge interactions over a wide range of temperatures, suggesting a mechanism of chromatin structural organization whereby histone tails regulate inter- and intranucleosomal assemblies via binding with nucleosomal DNA. We identify specific regions of the histone core H2A/H2B-H4/H3-H3/H4-H2B/H2A, termed "cold sites", which retain a significant fraction of contacts with adjoining residues throughout the simulation, indicating their functional role in nucleosome organization. Cold sites are clustered around H3-H3, H2A-H4 and H4-H2A interhistone interfaces, indicating the necessity of these contacts for nucleosome stability. Essential dynamics analysis of simulation trajectories shows that bending across the H3-H3 is a prominent mode of intranucleosomal dynamics. We postulate that effects of salts on mononucleosomes can be modeled in discrete molecular dynamics by modulating histone-DNA interaction potentials. Local fluctuations in nucleosomal DNA vary significantly along the DNA sequence, suggesting that only a fraction of histone-DNA contacts make strong interactions dominating mononucleosomal dynamics. Our findings suggest that histone tails have a direct functional role in stabilizing higher-order chromatin structure, mediated by salt-bridge interactions with adjacent DNA.

Chromatin↗

PubMed1998

Trajectory formation based on physiological characteristics of skeletal muscles.

Human arm trajectories in natural unrestricted reaching movements were studied. They have particular properties such that a hand path is a rather simple straight or curved line, and a tangential velocity profile of hand is bell-shaped. Also these properties are invariant, independent of movement duration and hand-held load. In this study, trajectory formation is investigated on the basis of physiological characteristics of skeletal muscles, and a criterion prescribed by a derivative of isometric muscle torque is proposed. Subsequently, optimal trajectories are formulated under various conditions of movement to account for a planning strategy of human arm trajectories. In addition to such a theoretical approach, human arm trajectories are experimentally observed by a measuring system which provides a visual sensor and a target tracking device, enabling totally unrestricted movements. Then, optimal trajectories are quantitatively evaluated in comparison with experimental data in which essential properties of human arm trajectories are demonstrated. These results support the idea that human arm trajectories are planned in order to minimize the proposed criterion which is determined from physiological aspects. Finally, the physiological advantages of human arm trajectories are discussed with regard to the analysis of observed and optimal trajectories.

Arm↗

PubMed2003

State and spectral properties of chloride oscillations in pollen.

Pollen tube growth is a dynamic system expressing a number of oscillating circuits. Our recent work identified a new circuit, oscillatory efflux of Cl(-) anion from the pollen tube apex. Cl(-) efflux is the first ion signal found to be coupled in phase with growth oscillations. Functional analyses indicate an active role for Cl(-) flux in pollen tube growth. In this report the dynamical properties of Cl(-) efflux are examined. Phase space analysis demonstrates that the system trajectory converges on a limit cycle. Fourier analysis reveals that two harmonic frequencies characterize normal growth. Cl(-) efflux is inhibited by the channel blocker DIDS, is stimulated by hypoosmotic treatment, and is antagonized by the signal encoded in inositol 3,4,5,6-tetrakisphosphate. These perturbations induce transitions of the limit cycle to new metastable states or cause system collapse to a static attractor centered near the origin. These perturbations also transform the spectral profile, inducing subharmonic frequencies, transitions to period doubling and tripling, superharmonic resonance, and chaos. These results indicate that Cl(-) signals in pollen tubes display features that are characteristic of active oscillators that carry frequency-encoded information. A reaction network of the Cl(-) oscillator coupled to two nonlinear feedback circuits that may drive pollen tube growth oscillations is considered.

4,4'-Diisothiocyanostilbene-2,2'-Disulfonic Acid↗

PubMed2006

Development and prediction of hyperactive symptoms from 2 to 7 years in a population-based sample.

OBJECTIVES: Children with hyperactive symptoms are often referred to mental health services. Given the frequency and persistent nature of hyperactivity, it is important to better understand its developmental course. This study identified the different developmental trajectories of hyperactive symptoms from 2 to 7 years and tested early predictors of high-level and persistent hyperactivity. These data may lead to earlier detection of at-risk children and to more effective interventions that take into account developmental considerations. PARTICIPANTS: Four data-collection cycles of a nationwide survey of Canadian children were used to track the early development of hyperactivity. Children were 0 to 23 months at the first cycle in 1994 and 6 to 7 years at the fourth cycle in 2000. OUTCOME MEASURES: Hyperactivity data were gathered from mothers on a biennial basis beginning when children were 24 months old. Information on potential prenatal and postnatal predictors was gathered from mothers at the first cycle. DESIGN: Group-based semiparametric mixture modeling was used to estimate developmental trajectories, and logistic-regression analysis identified predictors of hyperactivity. RESULTS: Four trajectories of hyperactive symptoms were identified: very low, low, moderate, and high. Statistically significant predictors for high and persistent hyperactivity, after controlling for all other factors, were maternal prenatal smoking, child male gender, maternal depression, and hostile parenting. CONCLUSIONS: For the majority of children, the frequency of hyperactive symptoms decreased or remained low from 2 to 7 years. However, 7 children in 100 were classified as having high initial levels of hyperactive symptoms that persisted over time. Several prenatal and early postnatal risk factors identified these children, although additional variables will need to be identified to accurately predict high and persistent hyperactivity. Findings suggest that preventive interventions could target high-risk families during pregnancy and early childhood.

Age of Onset↗

PubMed2004

Theoretical study of the ligand-CYP2B4 complexes: effect of structure on binding free energies and heme spin state.

The molecular origins of temperature-dependent ligand-binding affinities and ligand-induced heme spin state conversion have been investigated using free energy analysis and DFT calculations for substrates and inhibitors of cytochrome P450 2B4 (CYP2B4), employing models of CYP2B4 based on CYP2C5(3LVdH)/CYP2C9 crystal structures, and the results compared with experiment. DFT calculations indicate that large heme-ligand interactions (ca. -15 kcal/mol) are required for inducing a high to low spin heme transition, which is correlated with large molecular electrostatic potentials (approximately -45 kcal/mol) at the ligand heteroatom. While type II ligands often contain oxygen and nitrogen heteroatoms that ligate heme iron, DFT results indicate that BP and MF heme complexes, with weak substrate-heme interactions (ca. -2 kcal/mol), and modest MEPS minima (>-35 kcal/mol) are high spin. In contrast, heme complexes of the CYP2B4 inhibitor, 4PI, the product of benzphetamine metabolism, DMBP, and water are low spin, have substantial heme-ligand interaction energies (<-15 kcal/mol) and deep MEPS minima (<-45 kcal/mol) near their heteroatoms. MMPBSA analysis of MD trajectories were made to estimate binding free energies of these ligands at the heme binding site of CYP2B4. In order to initially assess the realism of this approach, the binding free energy of 4PI inhibitor was computed and found to be a reasonable agreement with experiment: -7.7 kcal/mol [-7.2 kcal/mol (experiment)]. BP was determined to be a good substrate [-6.3 kcal/mol (with heme-ligand water), -7.3 kcal/mol (without ligand water)/-5.8 kcal/mol (experiment)], whereas the binding of MF was negligible, with only marginal binding binding free energy of -1.7 kcal/mol with 2-MF bound [-3.8 kcal/mol (experiment)], both with and without retained heme-ligand water. Analysis of the free energy components reveal that hydrophobic/nonpolar contributions account for approximately 90% of the total binding free energy of these substrates and are the source of their differential and temperature-dependent CYP2B4 binding. The results indicate the underlying origins of the experimentally observed differential binding affinities of BP and MF, and indicate the plausibility of the use of models derived from moderate sequence identity templates in conjunction with approximate free energy methods in the estimation of ligand-P450 binding affinities.

Amino Acid Sequence↗

PubMed2003

Characteristics of trajectory in the migration of Amoeba proteus.

We investigated the behavior of migration of Amoeba proteus in an isotropic environment. We found that the trajectory in the migration of A. proteus is smooth in the observation time of 500-1000 s, but its migration every second (the cell velocity) on the trajectory randomly changes. Stochastic analysis of the cell velocity and the turn angle of the trajectory has shown that the histograms of the both variables well fit to Gaussian curves. Supposing a simple model equation for the cell motion, we have estimated the motive force of the migrating cell, which is of the order of piconewton. Furthermore, we have found that the cell velocity and the turn angle have a negative cross-correlation coefficient, which suggests that the amoeba explores better environment by changing frequently its migrating direction at a low speed and it moves rectilinearly to the best environment at a high speed. On the other hand, the model equation has simulated the negative correlation between the cell velocity and the turn angle. This indicates that the apparently rational behavior comes from intrinsic characteristics in the dynamical system where the motive force is not torquelike.

Amoeba↗

PubMedDate not supplied

Neuromuscular patterns of finger movements during piano playing. Definition of an experimental protocol.

Body movements require the activity of muscles fired by their motor neurons, controlled and coordinated according to central motor patterns modulated by the sensory feedback. The direct analysis of movements and forces produced by muscular activity can provide useful information into anatomical details of difficult observation. For instance, the analysis of movement trajectories can be used to obtain some insight into the underlying neuromuscular processes. Piano playing is a learned ability which links natural hand and finger movements in a complex and well-codified pattern. In the present study, the repeatability of finger movements was assessed in pianists of different experience. Five pianists (two females, 9, 23 years; three males, 24, 39, 70 years; all professionals) played nine two-octave C-major scales at 80, 112 and 160 bpm. The three-dimensional coordinates of their right hand fingers were digitized by a motion analyzer. For each pianist, data from each of the three repetition performed at the same velocity were aligned on the time of onset of the first C keypress (thumb), and a coefficient of superimposition was computed. Significant (analysis of variance, p < 0.001) differences were found between pianists and velocities, with a significant pianist x velocity interaction. The oldest man (a well-known concert player) was the less repeatable (mean superimposition 65.8%, SD 17), the 39-y-old man (a piano teacher) was the most repeatable (mean superimposition 81.4%, SD 8.1). In both of them, the thumb was the most repeatable finger; in the girl, the fifth and fourth fingers scored the best repeatability. The fastest scales were the less repeatable in the 70 and 24-y-old men, and the most repeatable in the 39-y-old man and 23-y-old woman. Even a simple and basic piano exercise can be performed with different movement patterns, without a direct relationship to the pianist experience. Apparently, repeatability was lower in concert pianists than in teachers and learners.

Adult↗

PubMed1997

Molecular dynamics simulations of the docking of substituted N5-deazapterins to dihydrofolate reductase.

Orientations of the deazapterin ring and the conformational preferences of groups appended to the deazapterin ring in a set of 8-substituted deazapterin cations docked into the dihydrofolate reductase (DHFR) binding site have been investigated using a methodology based on the simulated annealing technique within molecular dynamics (MD) simulations. Of five possible binding pockets for the 8-substituents, identified from a preliminary manual docking study, one has been definitively eliminated after an analysis of MD trajectories, while another remains uncertain. Using a new method based on standard thermodynamic cycles and a linear approximation of polar and non-polar free energy contributions from MD averages, binding affinities of the different ligands in each binding site have been correlated with experimental dissociation constants. The study has provided insights into structure-activity relationships for use in the design of modified inhibitors of DHFR.

Binding Sites↗

PubMed2004

Molecular dynamics simulations of glycosyltransferase LgtC.

Molecular dynamics simulations have been performed on fully solvated alpha-(1-->4)-galactosyltransferase LgtC from Neisseria meningitidis with and without the donor substrate UDP-Gal and in the presence of the manganese ion. The analysis of the trajectories revealed a limited movement in the loop X (residues 75-80) and a larger conformational change in the loop Y (residues 246-251) in the simulation, when UDP-Gal was not present. In this case, the loops X and Y open by almost 10A, exposing the active site to the solvent. The 'hinge region' responsible for the opening is composed of residues 246-247. We have also analyzed the behavior of the manganese ion in the simulations. The coordination number is 6 when UDP-Gal is present and it increases to 7 when it is absent. In the latter case, three water molecules become coordinated to the ion. In both cases, the coordination is very stable implying that the manganese ion is tightly bound in the active site of the enzyme even if UDP-Gal is not present. Further analysis of the structural water molecules location confirmed that the mobility of water molecules in the active site and the accessibility of this site for solvent are higher in the absence of the substrate.

Amino Acid Sequence↗

PubMed2001

Conformational space comparison of GnRH and lGnRH-III using molecular dynamics, cluster analysis and Monte Carlo thermodynamic integration.

The conformational space available to GnRH and lGnRH-III was compared using 5.2 ns constant temperature and pressure molecular dynamics simulations with explicit TIP3P solvation and the AMBER v. 5.0 force field. Cluster analysis of both trajectories resulted in two groups of conformations. Results of free energy calculations, in agreement with previous experimental data, indicate that a conformation with a turn from residues 5 through 8 is preferred for GnRH in an aqueous environment. By contrast, a conformation with a helix from residues 2 through 7 with a bend from residues 6 through 10 is preferred for lGnRH-III in an aqueous environment. The side chains of His2 and Trp3 in lGnRH-III occupy different regions of phase space and participate in weakly polar interactions different from those in GnRH. The unique conformational properties of lGnRH-III may account for its specific anti cancer activity.

Animals↗

PubMed2003

Straight-forward versus anatomic trajectory technique of thoracic pedicle screw fixation: a biomechanical analysis.

STUDY DESIGN: A biomechanical study on cadaveric thoracic vertebrae using pullout strength, insertional torque, and bone mineral density to determine the optimal sagittal trajectory of thoracic pedicle screws. OBJECTIVE: To perform a biomechanical study on cadaveric thoracic vertebrae using insertional torque, pullout strength, and bone mineral density to determine the optimal biomechanical sagittal trajectory for placement thoracic pedicle screws. We compared the straight-forward (paralleling the vertebral endplate) with anatomic trajectory (directed along the true anatomic axis of the pedicle). METHODS: Thirty cadaveric thoracic vertebrae were harvested and evaluated with dual-energy x-ray absorptiometry to assess bone mineral density. Matched, fixed-head pedicle screws were then randomly assigned by side and placed using the straight-forward or anatomic technique under fluoroscopic visualization while recording the maximum insertional torque. Pullout strength testing was then performed. RESULTS: The maximum insertional torque for the straight-forward technique was 2.58 +/- 0.14 (SE) in pounds, whereas the anatomic technique averaged 1.86 +/- 0.14 (SE) in pounds (P = 0.0005). The maximum insertional torque at the neurocentral junction for the straight-forward technique averaged 1.89 +/- 0.17 (SE) in-lbs. (73% of maximum insertional torque), whereas the anatomic trajectory averaged 1.39 +/- 0.11 (SE) in pounds (75% of maximum insertional torque) (P = 0.007). The average pullout strength using a straight-forward trajectory was 611 +/- 50 (SE) N compared to the anatomic trajectory, which averaged 481 +/- 54 (SE) N (P = 0.034). The pullout strength correlated with mean bone mineral density for both the straight-forward (r = 0.461, P = 0.027) and anatomic (r = 0.598, P = 0.004) techniques. CONCLUSIONS: The straight-forward technique results in a 39% increase in maximum insertional torque and a 27% increase in pullout strength compared to the anatomic technique. The maximum insertional torque at the neurocentral junction resulted in a 36% increase using the straight-forward technique versus the anatomic trajectory. Bone mineral density directly correlates with pullout strength for both techniques.

Absorptiometry, Photon↗

PubMed1998

Lattice models for proteins reveal multiple folding nuclei for nucleation-collapse mechanism.

The nature of the nucleation-collapse mechanism in protein folding is probed using 27-mer and 36-mer lattice models. Three different forms for the interaction potentials are used. Three of the four 27-mer sequences have maximally compact and identical native state while the other has a non-compact native conformation. All the sequences fold thermodynamically and kinetically by a two-state process. Analysis of individual trajectories for each sequence using a self-organizing neural net algorithm shows that upon formation of a critical set of contacts the polypeptide chain rapidly reaches the native conformation which is consistent with a nucleation-collapse mechanism. The algorithm, which reduces the identification of the folding nucleus for each trajectory to one of pattern recognition, is used to show that there are multiple folding nuclei. There is a distribution of nucleation contacts in the transition states with some of them occurring with more probability (when averaged over the denatured ensemble) than others. We also show that there is a distribution in the size of the nuclei with the average number of residues in the folding nuclei being less than about one-third of the chain size. The fluctuations in the sizes of the nuclei are large, suggestive of a broad transition region. The folding nuclei, the structures of each are the corresponding transition states, have varying degree of overlap with the native conformation. The distribution of the radius of gyration of the transition states shows that these structures are an expanded form (by about 25% in the radius of gyration) of the native conformation. Local contacts are most dominant in the folding nuclei while a certain fraction of non-local contacts is necessary to stabilize the transition states. The search for the critical nuclei initially involves the formation of local contacts, while non-local contacts are formed later. The fractional values of PhiF for the two 27-mer mutants found by using the protein engineering protocol are consistent with the microscopic picture of partial formation of structures involving these residues in the transition state. These observations lead to a multiple folding nuclei (MFN) model for nucleation-collapse mechanism in protein folding. The major implication of the MFN model is that, even if the residues whose tertiary interactions are formed nearly completely in the transition state are mutated, it does not disrupt the nature of the nucleation-collapse mechanism. We analyze the experiments on chymotrypsin inhibitor 2 and alpha-spectrin SH3 domain and two circular permutants in light of the MFN model. It is shown that the PhiF-value analysis for these proteins gives considerable support to the MFN model. The theoretical and experimental studies give a coherent picture of the nucleation-collapse mechanism in which there is a distribution of folding nuclei with some more probable than others. The formation of any specific nucleus is not necessary for efficient two-state folding.

Alanine↗

PubMed1997

Coping with systematic bias during bilateral movement.

The present studies examined the nature of kinematic interlimb interference during bilateral elbow movements of 1:1, 2:1 and 3:1 frequency ratios and the manner in which subjects cope with coordination bias. Analysis of movement trajectories in the first experiment indicated progressively greater angular velocity assimilation across 2:1 and 3:1 conditions. The desired temporal relationship was maintained by slowing or pausing the low-frequency movement at peak extension while the high-frequency arm produced intervening cycles. An increase in amplitude was also evident for concurrent, homologous cycles. Movement smoothness was emphasized and additional practice was provided in a second experiment. This resulted in dissociated peak angular velocity between limbs and eliminated hesitations and amplitude effects. Bias was still evident, however, as an intermittent approach toward a 1:1 ratio within each cycle. This systematic tendency was somewhat greater at the lower of two absolute frequency combinations but was not influenced by the role of each arm in producing the higher or lower frequency movement. The findings from the first experiment suggest that subjects initially accommodate interlimb kinematic assimilation, while producing the intended timing ratio, by intermittently slowing or pausing the lower-frequency movement. This attenuates the need for bilaterally-disparate movement parameters and provides additional time for organizing residual kinematic differences, perhaps reducing "transient coupling." Evidence from the second experiment indicates that subtle relative motion preferences are still evident following sufficient practice to perform the movements smoothly. The within-cycle locations of the points of greatest interlimb bias for the 2:1 rhythms were positively displaced from those previously observed for 1:1 oscillations. The persistent coordination tendencies noted in both experiments perhaps reflect an assimilation/compensation cycle and constitute one potential source of the systematic error that often emerges during the acquisition of complex skills.

Adult↗

PubMed2001

Role of salt bridges in homeodomains investigated by structural analyses and molecular dynamics simulations.

Homeodomains are a class of helix-turn-helix DNA-binding protein motifs that play an important role in the control of cellular development in eukaryotes. They fold in a three alpha-helix structural module, where the third helix is the recognition helix that fits into the major groove of DNA. Structural analysis of the members of the homeodomain family led to the identification of interactions likely to stabilize the protein domains. Linking the helices pairwise, three salt bridges were found to be well preserved within the family. Also well conserved were two cation-pi interactions between aromatic and positively charged side chains. To analyze the structural role of the salt bridges, molecular dynamics simulations (MD) were carried out on the wild-type homeodomain from the Drosophila paired protein (1fjl) and on three mutants, which lack one or two salt bridges and mimic natural mutations in other homeodomains. Analysis of the trajectories revealed only small structural rearrangements of the three helices in all MD simulations, thereby suggesting that the salt bridges have no essential stabilizing role at room temperature, but rather might be important for improving thermostability. The latter hypothesis is supported by a good correlation between the melting midpoint temperatures of several homeodomains and the number of salt bridges and cation-pi interactions that connect secondary structures.

Amino Acid Sequence↗

PubMed2000

Disruption of spatial organization and interjoint coordination in Parkinson's disease, progressive supranuclear palsy, and multiple system atrophy.

Patients with basal ganglia diseases may exhibit ideomotor apraxia. To define the nature of the impairment of the action production system, we studied a repetitive gesture of slicing bread by three-dimensional computergraphic analysis in eight nondemented patients with Parkinson's disease in the "on" state, five with progressive supranuclear palsy and four with multiple system atrophy. Two patients with Parkinson's disease and two with progressive supranuclear palsy showed ideomotor apraxia for transitive movements on standard testing. A Selspott II system was used for kinematic analysis of wrist trajectories and angular motions of the shoulder and elbow joints. Patients with Parkinson's disease, progressive supranuclear palsy, and even some with multiple system atrophy exhibited kinematic deficits in the spatial precision of movement and velocity-curvature relationships; in addition, they failed to maintain proper angle/angle relationships and to apportion their relative joint amplitudes normally. Spatial disruption of wrist trajectories was more severe in patients with ideomotor apraxia. We posit that the basal ganglia are part of the parallel parieto-frontal circuits devoted to sensorimotor integration for object-oriented behavior. The severity and characteristics of spatial abnormalities of a transitive movement would therefore depend on the location and distribution of the pathologic process within these circuits.

Aged↗

PubMed2005

Ontogeny of craniofacial sexual dimorphism in the orangutan (Pongo pygmaeus). I: face and palate.

The orangutan is widely recognized as a highly dimorphic species. An ontogenetic approach to the study of sexual dimorphism can assist researchers in understanding both where and when these differences develop. In this study, 357 orangutans from Borneo were divided into five developmental stages representing infancy to mature adulthood. Three-dimensional (3D) coordinate data from 16 landmarks representing the face and palate were analyzed by means of a Euclidean distance matrix analysis (EDMA), a quantitative method for the comparison of forms. Three separate analyses (an age-specific static comparison of forms, a sex-specific analysis of growth trajectories, and an intersex comparison of patterns of relative growth) were carried out with the intent to describe the rate, timing, magnitude, and pattern of growth in the orangutan face and palate. The results indicate that generally males and females share a similar, but not identical, pattern of growth or local form change, but differ in growth rate, timing, and magnitude of difference. Dimorphism in the face and palate can be localized in infancy and traced throughout all age intervals. Orangutan females grow slightly faster than males from infancy to adolescence, at which time male growth exceeds female growth. Female growth ceases with the advent of adulthood, while male growth continues (i.e., both the number and magnitude of the dimorphic dimensions increase). Males and females are similar in facial dimensions and growth related to the orbits, upper face, and palate width. They maintain these similarities throughout development. However, they differ in facial and nasal height, palate length, snout projection, depth of the nasopharynx, and hafting of the face onto the skull. The face broadens and the zygomatic bone flares dramatically in adult males, corresponding to the development of cheek pads. While growth patterns are similar between the two sexes, they differ in the lateral orbit, snout projection, and hafting of the face onto the cranium. Adult dimorphism is the result of growth patterns experienced throughout life, and it is not equally expressed across the cranium. An understanding of patterns of dimorphism, along with the magnitude of difference, may be helpful for interpreting dimorphism in the fossil record.

Animals↗