The face is not an empty canvas: how facial expressions interact with facial appearance

Faces are not simply blank canvases upon which facial expressions write their emotional messages. In fact, facial appearance and facial movement are both important social signalling systems in their own right. We here provide multiple lines of evidence for the notion that the social signals derived from facial appearance on the one hand and facial movement on the other interact in a complex manner, sometimes reinforcing and sometimes contradicting one another. Faces provide information on who a person is. Sex, age, ethnicity, personality and other characteristics that can define a person and the social group the person belongs to can all be derived from the face alone. The present article argues that faces interact with the perception of emotion expressions because this information informs a decoder''s expectations regarding an expresser''s probable emotional reactions. Facial appearance also interacts more directly with the interpretation of facial movement because some of the features that are used to derive personality or sex information are also features that closely resemble certain emotional expressions, thereby enhancing or diluting the perceived strength of particular expressions.     

Infectious disease and sickness behaviour: tumour progression affects interaction patterns and social network structure in wild Tasmanian devils

Infectious diseases, including transmissible cancers, can have a broad range of impacts on host behaviour, particularly in the latter stages of disease progression. However, the difficulty of early diagnoses makes the study of behavioural influences of disease in wild animals a challenging task. Tasmanian devils (Sarcophilus harrisii) are affected by a transmissible cancer, devil facial tumour disease (DFTD), in which tumours are externally visible as they progress. Using telemetry and mark–recapture datasets, we quantify the impacts of cancer progression on the behaviour of wild devils by assessing how interaction patterns within the social network of a population change with increasing tumour load. The progression of DFTD negatively influences devils'' likelihood of interaction within their network. Infected devils were more active within their network late in the mating season, a pattern with repercussions for DFTD transmission. Our study provides a rare opportunity to quantify and understand the behavioural feedbacks of disease in wildlife and how they may affect transmission and population dynamics in general.     

Predation risk and the evolutionary ecology of reproductive behaviour

A large literature exists on the effects of predation risk on foraging and survival-related behaviours. In contrast, with some notable exceptions, relatively few theoretical or experimental studies have examined the effects of predation risk on reproductive behaviours. Existing literature on risk and reproductive behaviour and suggestion directions for future study are reviewed. In particular: (1) effects on predation risk on mating behaviour; (2) the influence of spatial patchiness on interactions between risk and reproductive behaviour; (3) the potential influence of multi-species interactions on the effects of predation risk on mating dynamics; (4) the importance of looking at sets of inter-related antipredator traits; and (5) some effects of predation risk on prey population patterns due to changes in prey reproductive behaviour are discussed. To illustrate various points examples involving fish and other aquatic organisms are used.     

Variability,morphometrics, and co-variation of the os lacrimale in Cervidae

In Ruminantia, the lacrimal bone forms a considerable part of the facial skeleton, and the morphology of its facial facet is highly variable when compared to other mammals. In this study, we quantify the species-specific variability in size and shape of the lacrimal facial facet in species of Cervidae (deer) and relate it to systematics and various aspects of their ecology and behavior. We sampled 143 skull specimens from 10 genera; 12 Moschus and 3 Tragulus specimens were used as outgroups. We find that size and shape of the lacrimal facial facet allow differentiating most species analyzed here, except for Mazama gouazoubira and Capreolus capreolus. Size and shape of the lacrimal facial facet vary widely across Cervidae regardless of their systematic relationships, ecology or behavior. Thus, we could not detect a unique signature of adaptational criteria in lacrimal morphology. Our data indicate that the lacrimal facial facet scales allometrically with skull size, in particular, the lacrimojugal length scales positively and the lacrimomaxillar length scales negatively. However, correlation analyses did not reveal any differences in the integration of the lacrimal bone with any specific skull module in any of the species compared. Lastly, we could not ascertain any correlation between the size and position of the preorbital depression with the size and shape of the lacrimal facial facet. We conclude that the lacrimal facial facet is highly flexible and may rapidly adjust to its surrounding bones. Its allometric growth appears to be an example of exaptation: changes in size and shape in the context of the increase of the skull length provide lacrimal contacts, in particular, a lacrimojugal one, which may serve to reduce mechanical loads resulting from increasingly larger antlers in large cervids.     

Using robots to understand social behaviour

A major challenge in studying social behaviour stems from the need to disentangle the behaviour of each individual from the resulting collective. One way to overcome this problem is to construct a model of the behaviour of an individual, and observe whether combining many such individuals leads to the predicted outcome. This can be achieved by using robots. In this review we discuss the strengths and weaknesses of such an approach for studies of social behaviour. We find that robots—whether studied in groups of simulated or physical robots, or used to infiltrate and manipulate groups of living organisms—have important advantages over conventional individual‐based models and have contributed greatly to the study of social behaviour. In particular, robots have increased our understanding of self‐organization and the evolution of cooperative behaviour and communication. However, the resulting findings have not had the desired impact on the biological community. We suggest reasons for why this may be the case, and how the benefits of using robots can be maximized in future research on social behaviour.     

Facialmetric similarities mediate mate choice: sexual imprinting on opposite-sex parents

Former studies have suggested that imprinting-like processes influence the shaping of human mate preferences. In this study, we provide more direct evidence for assessing facial resemblance between subjects' partner and subjects' parents. Fourteen facial proportions were measured on 312 adults belonging to 52 families, and the correlations between family members were compared with those of pairs randomly selected from the population. Spouses proved to be assortatively mated in the majority of measured facial proportions. Significant correlations have been found between the young men and their partner's father (but not his mother), especially on facial proportions belonging to the central area of the face. Women also showed resemblance to their partner's mother (but not to their father) in the facial characteristics of their lower face. Replicating our previous studies, facial photographs of participants were also matched by independent judges who ascribed higher resemblance between partners, and subjects and their partners' opposite-sex parents, compared with controls. Our results support the sexual imprinting hypothesis which states that children shape a mental template of their opposite-sex parents and search for a partner who resembles that perceptual schema. The fact that only the facial metrics of opposite-sex parents showed resemblance to the partner's face tends to rule out the role of familiarity in shaping mating preferences. Our findings also reject several other rival hypotheses. The adaptive value of imprinting-related human mating is discussed, and a hypothesis is made of why different facial areas are involved in males' and females' search for resemblance.     

Personality influences the neural responses to viewing facial expressions of emotion

Cognitive research has long been aware of the relationship between individual differences in personality and performance on behavioural tasks. However, within the field of cognitive neuroscience, the way in which such differences manifest at a neural level has received relatively little attention. We review recent research addressing the relationship between personality traits and the neural response to viewing facial signals of emotion. In one section, we discuss work demonstrating the relationship between anxiety and the amygdala response to facial signals of threat. A second section considers research showing that individual differences in reward drive (behavioural activation system), a trait linked to aggression, influence the neural responsivity and connectivity between brain regions implicated in aggression when viewing facial signals of anger. Finally, we address recent criticisms of the correlational approach to fMRI analyses and conclude that when used appropriately, analyses examining the relationship between personality and brain activity provide a useful tool for understanding the neural basis of facial expression processing and emotion processing in general.     

Turning the other cheek: the viewpoint dependence of facial expression after-effects

How do we visually encode facial expressions? Is this done by viewpoint-dependent mechanisms representing facial expressions as two-dimensional templates or do we build more complex viewpoint independent three-dimensional representations? Recent facial adaptation techniques offer a powerful way to address these questions. Prolonged viewing of a stimulus (adaptation) changes the perception of subsequently viewed stimuli (an after-effect). Adaptation to a particular attribute is believed to target those neural mechanisms encoding that attribute. We gathered images of facial expressions taken simultaneously from five different viewpoints evenly spread from the three-quarter leftward to the three-quarter rightward facing view. We measured the strength of expression after-effects as a function of the difference between adaptation and test viewpoints. Our data show that, although there is a decrease in after-effect over test viewpoint, there remains a substantial after-effect when adapt and test are at differing three-quarter views. We take these results to indicate that neural systems encoding facial expressions contain a mixture of viewpoint-dependent and viewpoint-independent elements. This accords with evidence from single cell recording studies in macaque and is consonant with a view in which viewpoint-independent expression encoding arises from a combination of view-dependent expression-sensitive responses.     

Chimpanzee facial symmetry: a biometric measure of chimpanzee health

This paper reports a study of fitness indicator theory in chimpanzees. First, it establishes a theoretical perspective for the study of fitness indicator theory and the relationships among indicators of fitness in humans and other animals. Second, it describes a methodology for assessing facial fluctuating asymmetry (FA) in a sample (N = 21) of zoo chimpanzees (Pan troglodytes). Third, associations among chimpanzee facial FA and health are described. FA was positively associated with negative health symptoms, and negatively associated with general health. Results are discussed under the framework of good genes theory.     

Facial EMG Responses to Combat-Related Visual Stimuli in Veterans With and Without Posttraumatic Stress Disorder

Veterans with (n = 10) and without (n = 10) posttraumatic stress disorder (PTSD) participated in an exploratory study of facial reactivity to neutral slides and to slides depicting unpleasant combat-related material that were previously determined to be emotionally evocative. It was found that the zygomaticus major (cheek), masseter (jaw), and lateral frontalis (forehead) muscles were especially reactive to the combat slides in the veterans with PTSD, suggesting the importance of facial emotional expression in this disorder. The PTSD participants' self-reports of overall distress paralleled these effects. However, autonomic reactivity did not reflect general arousal effects due to the visual stimuli, showing both the sensitivity of facial muscle assessment in this context and the need for further research on the relationship between stimulus modality and physiological trauma reactions. Additional directions for research in this area are discussed including efforts to correlate subjective and physiological reactions.     

Adaptation and the genetics of social behaviour

In recent years much progress has been made towards understanding the selective forces involved in the evolution of social behaviour including conflicts over reproduction among group members. Here, I argue that an important additional step necessary for advancing our understanding of the resolution of potential conflicts within insect societies is to consider the genetics of the behaviours involved. First, I discuss how epigenetic modifications of behaviour may affect conflict resolution within groups. Second, I review known natural polymorphisms of social organization to demonstrate that a lack of consideration of the genetic mechanisms involved may lead to erroneous explanations of the adaptive significance of behaviour. Third, I suggest that, on the basis of recent genetic studies of sexual conflict in Drosophila, it is necessary to reconsider the possibility of within-group manipulation by means of chemical substances (i.e. pheromones). Fourth, I address the issue of direct versus indirect genetic effects, which is of particular importance for the study of behaviour in social groups. Fifth, I discuss the issue of how a genetic influence on dominance hierarchies and reproductive division of labour can have secondary effects, for example in the evolution of promiscuity. Finally, because the same sets of genes (e.g. those implicated in chemical signalling and the responses that are triggered) may be used even in species as divergent as ants, cooperative breeding birds and primates, an integration of genetic mechanisms into the field of social evolution may also provide unifying ideas.     

Three-dimensional human facial morphologies as robust aging markers

Aging is associated with many complex diseases. Reliable prediction of the aging process is important for assessing the risks of aging-associated diseases. However, despite intense research, so far there is no reliable aging marker. Here we addressed this problem by examining whether human 3D facial imaging features could be used as reliable aging markers. We collected > 300 3D human facial images and blood profiles well-distributed across ages of 17 to 77 years. By analyzing the morphological profiles, we generated the first comprehensive map of the aging human facial phenome. We identified quantitative facial features, such as eye slopes, highly associated with age. We constructed a robust age predictor and found that on average people of the same chronological age differ by ± 6 years in facial age, with the deviations increasing after age 40. Using this predictor, we identified slow and fast agers that are significantly supported by levels of health indicators. Despite a close relationship between facial morphological features and health indicators in the blood, facial features are more reliable aging biomarkers than blood profiles and can better reflect the general health status than chronological age.     

Facial width-to-height ratio predicts self-reported dominance and aggression in males and females,but a measure of masculinity does not

Recently, associations between facial structure and aggressive behaviour have been reported. Specifically, the facial width-to-height ratio (fWHR) is thought to link to aggression, although it is unclear whether this association is related to a specific dimension of aggression, or to a more generalized concept of dominance behaviour. Similarly, an association has been proposed between facial masculinity and dominant and aggressive behaviour, but, to date, this has not been formally tested. Because masculinity and fWHR are negatively correlated, it is unlikely that both signal similar behaviours. Here, we thus tested these associations and show that: (i) fWHR is related to both self-reported dominance and aggression; (ii) physical aggression, verbal aggression and anger, but not hostility are associated with fWHR; (iii) there is no evidence for a sex difference in associations between fWHR and aggression; and (iv) the facial masculinity index does not predict dominance or aggression. Taken together, these results indicate that fWHR, but not a measure of facial masculinity, cues dominance and specific types of aggression in both sexes.     

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.     

康复护理对面瘫患者的影响

目的探讨面瘫患者实施康复护理的效果。方法选择2013年1月~2015年12月我院神经科收治的面瘫患者80例为研究对象,随机分为对照组和观察组各40例,对照组给予常规护理,观察组在常规护理基础上实施康复护理,观察比较两组患者的治疗效果。结果观察组临床总有效率明显高于对照组,差异有统计学意义(P0.05)。结论面瘫患者实施康复护理可提高临床效果,促进面神经的恢复,值得临床应用。     

Adaptive walks on behavioural landscapes and the evolution of optimal behaviour by natural selection

Summary One of the main challenges to the adaptationist programme in general and to the use of optimality models in behavioural and evolutionary ecology in particular is that natural selection need not optimise fitness. This challenge is addressed by considering the evolution of optimal patch choice by natural selection. The behavioural model is based on a state variable approach in which a strategy consists of a sequence denoting the patch to be visited as a function of the organism's state and time. The optimal strategy maximises expected terminal reproduction. The fitnesses of alternative strategies are computed by iteration of the associated equations for fitness; this characterises the adaptive behavioural landscape. There may be enormous numbers of strategies that have near optimal fitnesses. A population model is used to connect frequencies of behavioural types from one generation to the next. Theories on adaptive walks on fitness landscapes are considered in the context of behaviour. The main result is that within the context of optimality arguments at selective equilibrium, sub-optimal behaviours can persist. General implications for research in behavioural ecology, including tests of behavioural theories, are discussed.     

Allometry of facial mobility in anthropoid primates: implications for the evolution of facial expression

Body size may be an important factor influencing the evolution of facial expression in anthropoid primates due to allometric constraints on the perception of facial movements. Given this hypothesis, I tested the prediction that observed facial mobility is positively correlated with body size in a comparative sample of nonhuman anthropoids. Facial mobility, or the variety of facial movements a species can produce, was estimated using a novel application of the Facial Action Coding System (FACS). I used FACS to estimate facial mobility in 12 nonhuman anthropoid species, based on video recordings of facial activity in zoo animals. Body mass data were taken from the literature. I used phylogenetic generalized least squares (PGLS) to perform a multiple regression analysis with facial mobility as the dependent variable and two independent variables: log body mass and dummy-coded infraorder. Together, body mass and infraorder explain 92% of the variance in facial mobility. However, the partial effect of body mass is much stronger than for infraorder. The results of my study suggest that allometry is an important constraint on the evolution of facial mobility, which may limit the complexity of facial expression in smaller species. More work is needed to clarify the perceptual bases of this allometric pattern.     

Cranial variation in prehistoric human skeletal remains from the Marianas

Nonmetric cranial variation and facial flatness of the Pacific and circum-Pacific populations are investigated. The peoples of the Marianas, eastern Polynesia and Hawaii form a cluster and show affinities in terms of nonmetric cranial variation with the Southeast and East Asians rather than with the Jomon-Ainu, a view which is widely supported by others. Facial flatness analysis also indicates that Polynesians have different patterns of facial prominence as compared with the Jomon-Ainu. These results increase the difficulty of accepting the Jomon-Pacific cluster proposed by Brace and his coworkers. Although genetic and nonmetric cranial variation reveal relatively close relationships, the Mariana skeletons are markedly different in facial flatness and limb bone morphology from those of Polynesians. Am J Phys Anthropol 104:399–410, 1997. © 1997 Wiley-Liss, Inc.     

Morphological integration of soft-tissue facial morphology in Down Syndrome and siblings

Down syndrome (DS), resulting from trisomy of chromosome 21, is the most common live-born human aneuploidy. The phenotypic expression of trisomy 21 produces variable, though characteristic, facial morphology. Although certain facial features have been documented quantitatively and qualitatively as characteristic of DS (e.g., epicanthic folds, macroglossia, and hypertelorism), all of these traits occur in other craniofacial conditions with an underlying genetic cause. We hypothesize that the typical DS face is integrated differently than the face of non-DS siblings, and that the pattern of morphological integration unique to individuals with DS will yield information about underlying developmental associations between facial regions. We statistically compared morphological integration patterns of immature DS faces (N = 53) with those of non-DS siblings (N = 54), aged 6-12 years using 31 distances estimated from 3D coordinate data representing 17 anthropometric landmarks recorded on 3D digital photographic images. Facial features are affected differentially in DS, as evidenced by statistically significant differences in integration both within and between facial regions. Our results suggest a differential affect of trisomy on facial prominences during craniofacial development.