Facial perception of conspecifics: chimpanzees (Pan troglodytes) preferentially attend to proper orientation and open eyes

This paper reports on the use of an eye-tracking technique to examine how chimpanzees look at facial photographs of conspecifics. Six chimpanzees viewed a sequence of pictures presented on a monitor while their eye movements were measured by an eye tracker. The pictures presented conspecific faces with open or closed eyes in an upright or inverted orientation in a frame. The results demonstrated that chimpanzees looked at the eyes, nose, and mouth more frequently than would be expected on the basis of random scanning of faces. More specifically, they looked at the eyes longer than they looked at the nose and mouth when photographs of upright faces with open eyes were presented, suggesting that particular attention to the eyes represents a spontaneous face-scanning strategy shared among monkeys, apes, and humans. In contrast to the results obtained for upright faces with open eyes, the viewing times for the eyes, nose, and mouth of inverted faces with open eyes did not differ from one another. The viewing times for the eyes, nose, and mouth of faces with closed eyes did not differ when faces with closed eyes were presented in either an upright or inverted orientation. These results suggest the possibility that open eyes play an important role in the configural processing of faces and that chimpanzees perceive and process open and closed eyes differently.     

Perceptual integrality of componential and configural information in faces

The relative contribution of componential and configural information to face perception is controversial. We addressed this issue in the present study by examining how componential information and configural information interact during face processing, using Garner’s (1974) speeded classification paradigm. When classifying upright faces varying in components (eyes, nose, and mouth) and configural information (intereyes and nose-mouth spacing), observers could not selectively attend to components without being influenced by irrelevant variation in configural information, and vice versa, indicating that componential information and configural information are integral in upright face processing. Performance with inverted faces showed selective attention to components but not to configural information, implying dominance of componential information in processing inverted faces. When faces varied only in components, selective attention to different components was observed in upright and inverted faces, indicating that facial components are perceptually separable. These results provide strong evidence that integrality of componential and configural information, rather than the relative dominance of either, is the hallmark of upright face perception.     

Revisiting the perception of upside-down faces

In two experiments, the effect of orientation on face perception was assessed. Using a scale from 1 ( normal ) to 7 ( bizarre ), participants rated normal, unaltered faces and faces in which changes had been made to spatial-relational properties (eyes and mouth inverted or relative position of the eyes and mouth altered) or to component properties (eyes whitened and teeth blackened). For unaltered and component-distortion faces, bizarreness ratings increased linearly as orientation increased from 0° to 180°. For spatial-distortion faces, a discontinuity in the function relating orientation and bizarreness was in evidence between 90° and 120°. The results provide support for the view that there is a qualitative difference in the processing of upright and inverted faces due to the disproportionate effect of inversion on the encoding of spatial-relational information.     

A unique look at face processing: the impact of masked faces on the processing of facial features

This experiment utilized a masked priming paradigm to explore the early processes involved in face recognition. The first experiment investigated implicit processing of the eyes and mouth in an upright face, using prime durations of 33 and 50 ms. The results demonstrate implicit processing of both the eyes and mouth, and support the configural processing theory of face processing. The second experiment used the same method with inverted faces and the third experiment was a combination of Experiments 1 and 2. The fourth experiment utilized misaligned faces as the primes. Based on the pattern of results from these experiments, we suggest that, when a face is inverted, the eyes and mouth are initially processed individually and are not linked until a later stage of processing. An upright face is proposed to be processed by analysis of its configuration, whereas an inverted face is initially processed using first-order relational information, and then converted to an upright representation and transferred to face specific regions for configural analysis.     

Losing face: impaired discrimination of featural and configural information in the mouth region of an inverted face

Given that all faces share the same set of features—two eyes, a nose, and a mouth—that are arranged in similar configuration, recognition of a specific face must depend on our ability to discern subtle differences in its featural and configural properties. An enduring question in the face-processing literature is whether featural or configural information plays a larger role in the recognition process. To address this question, the face dimensions task was designed, in which the featural and configural properties in the upper (eye) and lower (mouth) regions of a face were parametrically and independently manipulated. In a same–different task, two faces were sequentially presented and tested in their upright or in their inverted orientation. Inversion disrupted the perception of featural size (Exp. 1), featural shape (Exp. 2), and configural changes in the mouth region, but it had relatively little effect on the discrimination of featural size and shape and configural differences in the eye region. Inversion had little effect on the perception of information in the top and bottom halves of houses (Exp. 3), suggesting that the lower-half impairment was specific to faces. Spatial cueing to the mouth region eliminated the inversion effect (Exp. 4), suggesting that participants have a bias to attend to the eye region of an inverted face. The collective findings from these experiments suggest that inversion does not differentially impair featural or configural face perceptions, but rather impairs the perception of information in the mouth region of the face.     

Attention to faces: a change-blindness study

What strategies does human vision use to attend to faces and their features? How are such strategies altered by 2-D inversion or photographic negation? We report two experiments in which these questions were studied with the flicker task of the change-blindness literature. In experiment 1 we studied detection of configural changes to the eyes or mouth, and found that upright faces receive more efficient attention than inverted faces, and that faces shown with normal contrast receive more efficient attention than faces shown in photographic negative. Moreover, eyes receive greater attention than the mouth. In experiment 2 we studied detection of local changes to the eyes or mouth, and found the same results. It is well known that inversion and negation impair the perception and recognition of faces. The experiments presented here extend previous findings by showing that inversion and negation also impair attention to faces.     

知觉场理论对面孔知觉年龄偏差效应的解释

使用融合面孔范式和倒置面孔范式来研究面孔知觉的年龄偏差效应,检验知觉场能否作为面孔整体加工的指标,并以此发展年龄偏差的整体加工解释。结果发现：（1）成人和儿童在加工正立面孔时的知觉场均大于加工倒置面孔时的知觉场;（2）在加工正立面孔时,成人加工本年龄面孔比加工他年龄面孔的知觉场更大。上述结果表明：（1）知觉场大小可以作为面孔整体加工的指标,且受面孔朝向的影响;（2）知觉场假设可以解释面孔的年龄偏差效应。     

知觉场理论对面孔知觉年龄偏差效应的解释

使用融合面孔范式和倒置面孔范式来研究面孔知觉的年龄偏差效应,检验知觉场能否作为面孔整体加工的指标,并以此发展年龄偏差的整体加工解释。结果发现：（1）成人和儿童在加工正立面孔时的知觉场均大于加工倒置面孔时的知觉场;（2）在加工正立面孔时,成人加工本年龄面孔比加工他年龄面孔的知觉场更大。上述结果表明：（1）知觉场大小可以作为面孔整体加工的指标,且受面孔朝向的影响;（2）知觉场假设可以解释面孔的年龄偏差效应。     

Automatic gaze to the nose region cannot be inhibited during observation of facial expression in Eastern observers

Humans can extract a great deal of information about others very quickly. This is partly because the face automatically captures observers’ attention. Specifically, the eyes can attract overt attention. Although it has been reported that not only the eyes but also the nose can capture initial oculomotor movement in Eastern observers, its generalizability remains unknown. In this study, we applied the “don’t look” paradigm wherein participants are asked not to fixate on a specific facial region (i.e., eyes, nose, and mouth) during an emotion recognition task with upright (Experiment 1) and inverted (Experiment 2) faces. In both experiments, we found that participants were less able to inhibit the initial part of their fixations to the nose, which can be interpreted as the nose automatically capturing attention. Along with previous studies, our overt attention tends to be attracted by a part of the face, which is the nose region in Easterner observers.     

Effects of motion and audio-visual redundancy on upright and inverted face and feature preferences in 4-13-month old pre- and full-term NICU graduates

NICU infants are reported to have diminished social orientation and increased risk of socio-communicative disorders. In this eye tracking study, we used a preference for upright compared to inverted faces as a gauge of social interest in high medical risk full- and pre-term NICU infants. We examined the effects of facial motion and audio-visual redundancy on face and eye/mouth preferences across the first year. Upright and inverted baby faces were simultaneously presented in a paired-preference paradigm with motion and synchronized vocalization varied. NICU risk factors including birth weight, sex, and degree of CNS injury were examined. Overall, infants preferred the more socially salient upright faces, making this the first report, to our knowledge, of an upright compared to inverted face preference among high medical risk NICU infants. Infants with abnormalities on cranial ultrasound displayed lower social interest, i.e. less of a preferential interest in upright faces, when viewing static faces. However, motion selectively increased their upright face looking time to a level equal that of infants in other CNS injury groups. We also observed an age-related sex effect suggesting higher risk in NICU males. Females increased their attention to the mouth in upright faces across the first year, especially between 7–10 months, but males did not. Although vocalization increased diffuse attention toward the screen, contrary to our predictions, there was no evidence that the audio-visual redundancy embodied in a vocalizing face focused additional attention on upright faces or mouths. This unexpected result may suggest a vulnerability in response to talking faces among NICU infants that could potentially affect later verbal and socio-communicative development.     

The ups and downs of face perception: evidence for holistic encoding of upright and inverted faces

A visual-search task was used to investigate the influence of facial organisation on discrimination of an internal facial feature. Participants searched for a downturned mouth in arrays of one to six faces that differed only in the target feature, with distractor faces containing an upturned mouth. Feature search was tested in four different face contexts: upright unaltered faces, inverted unaltered faces, upright faces in which the internal features were scrambled, or inverted scrambled faces. Normal face organisation facilitated feature search in upright faces, but slowed it in inverted faces. These findings demonstrate an interdependence of features and their configuration in the perceptual analysis of both upright and inverted faces.     

Left gaze bias in humans, rhesus monkeys and domestic dogs

While viewing faces, human adults often demonstrate a natural gaze bias towards the left visual field, that is, the right side of the viewee’s face is often inspected first and for longer periods. Using a preferential looking paradigm, we demonstrate that this bias is neither uniquely human nor limited to primates, and provide evidence to help elucidate its biological function within a broader social cognitive framework. We observed that 6-month-old infants showed a wider tendency for left gaze preference towards objects and faces of different species and orientation, while in adults the bias appears only towards upright human faces. Rhesus monkeys showed a left gaze bias towards upright human and monkey faces, but not towards inverted faces. Domestic dogs, however, only demonstrated a left gaze bias towards human faces, but not towards monkey or dog faces, nor to inanimate object images. Our findings suggest that face- and species-sensitive gaze asymmetry is more widespread in the animal kingdom than previously recognised, is not constrained by attentional or scanning bias, and could be shaped by experience to develop adaptive behavioural significance.     

When feature information comes first! Early processing of inverted faces

We investigated the early stages of face recognition and the role of featural and holistic face information. We exploited the fact that, on inversion, the alienating disorientation of the eyes and mouth in thatcherised faces is hardly detectable. This effect allows featural and holistic information to be dissociated and was used to test specific face-processing hypotheses. In inverted thatcherised faces, the cardinal features are already correctly oriented, whereas in undistorted faces, the whole Gestalt is coherent but all information is disoriented. Experiment 1 and experiment 3 revealed that, for inverted faces, featural information processing precedes holistic information. Moreover, the processing of contextual information is necessary to process local featural information within a short presentation time (26 ms). Furthermore, for upright faces, holistic information seems to be available faster than for inverted faces (experiment 2). These differences in processing inverted and upright faces presumably cause the differential importance of featural and holistic information for inverted and upright faces.     

Face and mouth inversion effects on visual and audiovisual speech perception

Three experiments examined whether image manipulations known to disrupt face perception also disrupt visual speech perception. Research has shown that an upright face with an inverted mouth looks strikingly grotesque whereas an inverted face and an inverted face containing an upright mouth look relatively normal. The current study examined whether a similar sensitivity to upright facial context plays a role in visual speech perception. Visual and audiovisual syllable identification tasks were tested under 4 presentation conditions: upright face-upright mouth, inverted face-inverted mouth, inverted face-upright mouth, and upright face-inverted mouth. Results revealed that for some visual syllables only the upright face-inverted mouth image disrupted identification. These results suggest that upright facial context can play a role in visual speech perception. A follow-up experiment testing isolated mouths supported this conclusion.     

First fixations in face processing: The more diagnostic they are the smaller the face-inversion effect

Hills, Ross, and Lewis (2011) introduced the concept that the face-inversion effect may, in part, be carried by the first feature attended to, since the first feature fixated upon is different for upright and inverted faces. An eye-tracking study that directly assesses this hypothesis by using fixation crosses to guide attention to the eye or mouth region of the to-be-presented upright and inverted faces was devised. Recognition was better when the fixation cross appeared at the eye region than at the mouth region. The face-inversion effect was smaller when the eyes were cued than when the mouth was cued or when there was no cueing. The eye-tracking measures confirmed that the fixation crosses attracted the first fixation but did not affect other measures of eye-movements. Furthermore, the location of the first fixation predicted recognition accuracy: when the first fixation was to the eyes, recognition accuracy was higher than when the first fixation was to the mouth, irrespective of facial orientation. The results suggest that the first facial feature attended to is more predictive of recognition accuracy than the face orientation in which they are presented.     

Early detection and avoidance of threatening faces during passive viewing

To evaluate whether there is an early attentional bias towards negative stimuli, we tracked participants' eyes while they passively viewed displays composed of four Ekman faces. In Experiment 1 each display consisted of three neutral faces and one face depicting fear or happiness. In half of the trials, all faces were inverted. Although the passive viewing task should have been very sensitive to attentional biases, we found no evidence that overt attention was biased towards fearful faces. Instead, people tended to actively avoid looking at the fearful face. This avoidance was evident very early in scene viewing, suggesting that the threat associated with the faces was evaluated rapidly. Experiment 2 replicated this effect and extended it to angry faces. In sum, our data suggest that negative facial expressions are rapidly analysed and influence visual scanning, but, rather than attract attention, such faces are actively avoided.     

How the Thatcher illusion reveals evolutionary differences in the face processing of primates

Face recognition in humans is a complex cognitive skill that requires sensitivity to unique configurations of eyes, mouth, and other facial features. The Thatcher illusion has been used to demonstrate the importance of orientation when processing configural information within faces. Transforming an upright face so that the eyes and mouth are inverted renders the face grotesque; however, when this “Thatcherized” face is inverted, the effect disappears. Due to the use of primate models in social cognition research, it is important to determine the extent to which specialized cognitive functions like face processing occur across species. To date, the Thatcher illusion has been explored in only a few species with mixed results. Here, we used computerized tasks to examine whether nonhuman primates perceive the Thatcher illusion. Chimpanzees and rhesus monkeys were required to discriminate between Thatcherized and unaltered faces presented upright and inverted. Our results confirm that chimpanzees perceived the Thatcher illusion, but rhesus monkeys did not, suggesting species differences in the importance of configural information in face processing. Three further experiments were conducted to understand why our results differed from previously published accounts of the Thatcher illusion in rhesus monkeys.     

Attention misplaced: the role of diagnostic features in the face-inversion effect

Inversion disproportionately impairs recognition of face stimuli compared to nonface stimuli arguably due to the holistic manner in which faces are processed. A qualification is put forward in which the first point fixated on is different for upright and inverted faces and this carries some of the face-inversion effect. Three experiments explored this possibility by using fixation crosses to guide attention to the eye or mouth region of the to-be-presented faces in different orientations. Recognition was better when the fixation cross appeared at the eye region than at the mouth region. The face-inversion effect was smaller when the eyes were cued than when the mouth was cued or when there was no cueing. The results suggest that the first facial feature attended to is important for accurate face recognition and this may carry some of the effects of inversion.     

The representations of spacing and part-based information are associated for upright faces but dissociated for objects: Evidence from individual differences

Considerable evidence suggests that qualitatively different processes are involved in the perception of faces and objects. According to a central hypothesis, the extraction of information about the spacing among face parts (e.g., eyes and mouth) is a primary function of face processing mechanisms that is dissociated from the extraction of information about the shape of these parts. Here, we used an individual-differences approach to test whether the shape of face parts and the spacing among them are indeed processed by dissociated mechanisms. To determine whether the pattern of findings that we reveal is unique for upright faces, we also presented similarly manipulated nonface stimuli. Subjects discriminated upright or inverted faces or houses that differed in parts or spacing. Only upright faces yielded a large positive correlation across subjects between performance on the spacing and part discrimination tasks. We found no such correlation for inverted faces or houses. Our findings suggest that face parts and spacing are processed by associated mechanisms, whereas the parts and spacing of nonface objects are processed by distinct mechanisms. These results may be consistent with the idea that faces are special, in that they are processed as nondecomposable wholes.     

眼睛区域构型信息与特征信息的跨维共变增益效应及其加工特异性

面孔知觉可能在区域尺度上发生多维信息整合, 但迄今无特异性实验证据。本研究在两个实验中操纵面孔眼睛区域或嘴巴区域的单维构型或特征信息, 测量人们觉察单维变化或跨维共变的敏感度, 以此检测面孔区域尺度上的多维信息整合有何现象与规律, 进而揭示面孔知觉的多维信息整合机制。实验获得3个发现：(1)正立面孔眼睛区域的信息变化觉察呈现出“跨维共变增益效应”, 跨维信息共变觉察的敏感度显著高于任意一种单维信息变化觉察的敏感度; (2)“跨维共变增益效应”只在正立面孔的眼睛区域出现, 在倒置面孔的眼睛区域、正立面孔的嘴巴区域或倒置面孔的嘴巴区域都没有出现, 因此具有面孔区域特异性和面孔朝向特异性; (3)就单维信息变化觉察而言, 眼睛区域的敏感度不会受到面孔倒置的损伤, 但是嘴巴区域的敏感度会受到面孔倒置的显著损伤。综合可知, 面孔知觉确实会发生区域尺度上的信息整合, 它不是普遍性的信息量效应, 而是特异性的眼睛效应(只发生在正立面孔的眼睛区域)。这是面孔整体加工(face holistic processing)在单维信息分辨和多维信息整合之间建立联系的必经环节。这提示我们对全脸多维信息知觉整合的理解需要从传统的面孔整体加工假设升级到以眼睛为中心的层级化多维信息整合算法(a hierarchical algorithm for multi-dimensional information integration)。