Discrimination of human and dog faces and inversion responses in domestic dogs (Canis familiaris)

Although domestic dogs can respond to many facial cues displayed by other dogs and humans, it remains unclear whether they can differentiate individual dogs or humans based on facial cues alone and, if so, whether they would demonstrate the face inversion effect, a behavioural hallmark commonly used in primates to differentiate face processing from object processing. In this study, we first established the applicability of the visual paired comparison (VPC or preferential looking) procedure for dogs using a simple object discrimination task with 2D pictures. The animals demonstrated a clear looking preference for novel objects when simultaneously presented with prior-exposed familiar objects. We then adopted this VPC procedure to assess their face discrimination and inversion responses. Dogs showed a deviation from random behaviour, indicating discrimination capability when inspecting upright dog faces, human faces and object images; but the pattern of viewing preference was dependent upon image category. They directed longer viewing time at novel (vs. familiar) human faces and objects, but not at dog faces, instead, a longer viewing time at familiar (vs. novel) dog faces was observed. No significant looking preference was detected for inverted images regardless of image category. Our results indicate that domestic dogs can use facial cues alone to differentiate individual dogs and humans and that they exhibit a non-specific inversion response. In addition, the discrimination response by dogs of human and dog faces appears to differ with the type of face involved.     

Face processing and familiarity: Evidence from eye‐movement data

How is information extracted from familiar and unfamiliar faces? Three experiments, in which eye‐movement measures were used, examined whether there was differential sampling of the internal face region according to familiarity. Experiment 1 used a face familiarity task and found that whilst the majority of fixations fell within the internal region, there were no differences in the sampling of this region according to familiarity. Experiment 2 replicated these findings, using a standard recognition memory paradigm. The third experiment employed a matching task, and once again found that the majority of fixations fell within the internal region. Additionally, this experiment found that there was more sampling of the internal region when faces were familiar compared with when they were unfamiliar. The use of eye fixation measures affirms the importance of internal facial features in the recognition of familiar faces compared with unfamiliar faces, but only when viewers compare pairs of faces.     

Aberrant first fixations when looking at inverted faces in various poses: The result of the centre‐of‐gravity effect?

Face recognition is essential in everyday human life, and all faces are encountered in different poses. However, when a face is inverted, difficulties arise for recognition and eye movements may (Barton, Radcliffe, Cherkasova, Edleman, & Intriligator, 2006) or may not be disrupted (Williams & Henderson, 2007). The present study explored the effects of orientation and pose on recognition and eye movements during a standard old/new recognition task in order to resolve whether inversion disrupts eye movements. Eye‐tracking data looked at the first fixations, the number of fixations, and the duration of fixations over a face. A standard inversion effect was observed, but the three‐quarter view advantage was not observed. Eye‐movement data revealed that the eyes were the most sampled feature (in terms of first fixation, number of fixations, and duration of fixation) for all upright faces, however, other features were sampled first for inverted faces. These results are consistent with Barton et al.'s (2006) but not Williams and Henderson's (2007) findings: possible explanations for this are discussed with the caveat that the same images were used from learning to test.     

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)。     

Self-face recognition is characterized by “bilateral gain” and by faster,more accurate performance which persists when faces are inverted

We examine interhemispheric cooperation in the recognition of personally known faces whose long-term familiarity ensures frequent co-activation of face-sensitive areas in the right and left brain. Images of self, friend, and stranger faces were presented for 150 ms in upright and inverted orientations both unilaterally, in the right or left visual field, and bilaterally. Consistent with previous research, we find a bilateral advantage for familiar but not for unfamiliar faces, and we demonstrate that this gain occurs for inverted as well as upright faces. We show that friend faces are recognized more quickly than unfamiliar faces in upright but not in inverted orientations, suggesting that configural processing underlies this particular advantage. Novel to this study is the finding that people are faster and more accurate at recognizing their own face over both stranger and friend faces and that these advantages occur for both upright and inverted faces. These findings are consistent with evidence for a bilateral representation of self-faces.     

Beyond U-Shaped Development in Infants' Processing of Faces: An Information-Processing Account

The development of the "inversion" effect in face processing was examined in infants 3 to 6 months of age by testing their integration of the internal and external features of upright and inverted faces using a variation of the "switch" visual habituation paradigm. When combined with previous findings showing that 7-month-olds use integrative processing of an upright face, but featural processing of an inverted face (Cohen & Cashon, 2001a), the present findings suggest that from 3 to 7 months, infants' ability to integrate facial features follows an N-shaped developmental pattern for upright faces and an inverted U-shaped pattern for inverted faces. We discuss these results in terms of a set of domain-general information-processing principles.     

Categorical perception effects for facial identity in robustly represented familiar and self-faces: the role of configural and featural information

Categorical perception of robustly represented faces (self, friend) and unfamiliar faces is investigated, and the relative roles of configural and featural information are examined. Participants performed identification and discrimination tasks on morph series containing the self-face and a friend's face (self-Friend 1), two friends' faces (Friend 2-Friend 3), and two unfamiliar faces (Unfamiliar 1-Unfamiliar 2), presented in upright and inverted orientations. For upright faces, categorical perception effects were observed for both familiar morph series but not for the unfamiliar morph series, suggesting that robust representation is a requirement for categorical perception in facial identity. For inverted faces, categorical perception was observed for the self-Friend 1 morph series only. This suggests that categorical perception is tied to configural processing for familiar non-self-faces, but can be observed for self-faces during featural processing-consistent with evidence that self-face representations contain strong configural and featural components. Finally, categorical perception is not enhanced by the presence of the self-face relative to other familiar faces when upright, but shows a trend of being enhanced for self-faces when inverted, adding to the debate on the ways in which robustly represented faces can elicit categorical perception.     

How do rhesus monkeys (<Emphasis Type="Italic">Macaca mulatta</Emphasis>) scan faces in a visual paired comparison task?

When novel and familiar faces are viewed simultaneously, humans and monkeys show a preference for looking at the novel face. The facial features attended to in familiar and novel faces, were determined by analyzing the visual exploration patterns, or scanpaths, of four monkeys performing a visual paired comparison task. In this task, the viewer was first familiarized with an image and then it was presented simultaneously with a novel and the familiar image. A looking preference for the novel image indicated that the viewer recognized the familiar image and hence differentiates between the familiar and the novel images. Scanpaths and relative looking preference were compared for four types of images: (1) familiar and novel objects, (2) familiar and novel monkey faces with neutral expressions, (3) familiar and novel inverted monkey faces, and (4) faces from the same monkey with different facial expressions. Looking time was significantly longer for the novel face, whether it was neutral, expressing an emotion, or inverted. Monkeys did not show a preference, or an aversion, for looking at aggressive or affiliative facial expressions. The analysis of scanpaths indicated that the eyes were the most explored facial feature in all faces. When faces expressed emotions such as a fear grimace, then monkeys scanned features of the face, which contributed to the uniqueness of the expression. Inverted facial images were scanned similarly to upright images. Precise measurement of eye movements during the visual paired comparison task, allowed a novel and more quantitative assessment of the perceptual processes involved the spontaneous visual exploration of faces and facial expressions. These studies indicate that non-human primates carry out the visual analysis of complex images such as faces in a characteristic and quantifiable manner.     

Internal feature saliency as a marker of familiarity and configural processing

Two experiments that explore the internal feature advantage (IFA) in familiar face processing are reported. The IFA involves more efficient processing of internal features for familiar faces over unfamiliar ones. Experiment 1 examined the possibility of a configural basis for this effect through use of a matching task for familiar and unfamiliar faces presented both upright and upside-down. Results revealed the predicted IFA for familiar faces when stimuli were upright, but this was removed when stimuli were inverted. Experiment 2 examined the degree of training required before the IFA was demonstrated. Latency results revealed that whilst 90–180 s of exposure was sufficient to generate an IFA of intermediate magnitude, 180–270 s of exposure was required before the IFA was equivalent to that demonstrated for a familiar face. Taken together, these results offer three conclusions: First, the IFA is reaffirmed as an objective indicator of familiarity; second, the IFA is seen to rest on configural processing; and finally, the development of the IFA with familiarity indicates a development of configural processing with familiarity. As such, insight is gained as to the type of processing changes that occur as familiarity is gradually acquired.     

Face processing in humans and new world monkeys: the influence of experiential and ecological factors

This study tests whether the face-processing system of humans and a nonhuman primate species share characteristics that would allow for early and quick processing of socially salient stimuli: a sensitivity toward conspecific faces, a sensitivity toward highly practiced face stimuli, and an ability to generalize changes in the face that do not suggest a new identity, such as a face differently oriented. The look rates by adult tamarins and humans toward conspecific and other primate faces were examined to determine if these characteristics are shared. A visual paired comparison (VPC) task presented subjects with either a human face, chimpanzee face, tamarin face, or an object as a sample, and then a pair containing the previous stimulus and a novel stimulus was presented. The stimuli were either presented all in an upright orientation, or all in an inverted orientation. The novel stimulus in the pair was either an orientation change of the same face/object or a new example of the same type of face/object, and the stimuli were shown either in an upright orientation or in an inverted orientation. Preference to novelty scores revealed that humans attended most to novel individual human faces, and this effect decreased significantly if the stimuli were inverted. Tamarins showed preferential looking toward novel orientations of previously seen tamarin faces in the upright orientation, but not in an inverted orientation. Similarly, their preference to look longer at novel tamarin and human faces within the pair was reduced significantly with inverted stimuli. The results confirmed prior findings in humans that novel human faces generate more attention in the upright than in the inverted orientation. The monkeys also attended more to faces of conspecifics, but showed an inversion effect to orientation change in tamarin faces and to identity changes in tamarin and human faces. The results indicate configural processing restricted to particular kinds of primate faces by a New World monkey species, with configural processing influenced by life experience (human faces and tamarin faces) and specialized to process orientation changes specific to conspecific faces.     

面孔倒置效应的研究与理论述评

面孔倒置效应是指, 个体对倒置面孔的再认成绩显著低于对正立面孔的再认成绩, 而且, 与普通物体的倒置效应相比, 面孔的倒置效应更大。研究者们针对面孔倒置效应提出了多种理论假说, 两种主要的理论解释是：第一, 倒置面孔破坏了面孔的结构特征, 影响了面孔的整体加工, 所以, 个体对倒置面孔的再认成绩较差; 第二, 眼睛在面孔加工中起重要作用, 尤其在倒置面孔中的作用更明显, 使得倒置面孔N170波幅较高。ERP和fMRI研究提示, 未来研究需要进一步明确眼睛、注意等在面孔倒置效应中的作用。     

On facial expertise: processing strategies of twins' parents

Parents of monozygotic twins typically learn to recognise their own children and also to tell them apart on photographs. However, it is unknown whether such exemplar expertise can be generalised to unfamiliar faces of equal similarity (ie other twins). In the present study, parents of monozygotic twins were compared to 'control' parents whose children were non-twin siblings. In a same/different task with familiar and unfamiliar twin faces as stimuli, twins' parents were faster than control parents in distinguishing their own children, but slower when distinguishing unfamiliar twin faces. These unexpected findings can be interpreted as a generalisation of a habitual perceptual strategy that is efficient only with familiar face exemplars. When pictures of the twins' faces were inverted by 180degrees, the typical inversion costs for familiar faces were observed, but the twins' parents recognised unfamiliar twins' faces better when these were inverted than when upright. This inverted inversion effect could be caused by the high similarity of the face pairs as well as by experience with familiar faces of equally high similarity. We conclude that the facial expertise of twins' parents is idiosyncratic to their own children and not based on an enhanced general facial expertise.     

Information processing during face recognition: the effects of familiarity, inversion, and morphing on scanning fixations

Where we make ocular fixations when viewing an object likely reflects interactions between 'external' object properties and internal 'top-down' factors, as our perceptual system tests hypotheses and attempts to make decisions about our environment. These scanning fixation patterns can tell us how and where the visual system gathers information critical to specific tasks. We determined the effects of the internal factors of expertise, experience, and ambiguity on scanning during a face-recognition task, in eight subjects. To assess the effects of expertise, we compared upright with inverted faces, since it is hypothesized that inverted faces do not access an orientation-dependent face-expert processor. To assess the effects of experience, we compared famous with novel faces, as famous faces would have stronger internal representations than anonymous ones. Ambiguity in matching seen and remembered faces was manipulated with morphed faces. We measured three classes of variables: (i) total scanning time and fixations; (ii) the spatial distribution of scanning; and (iii) the sequence of scanning, using first-order Markov matrices for local scan structure and string editing for global scan structure. We found that, with inverted faces, subjects redistributed fixations to the mouth and lower face, and their local and global scan structure became more random. With novel or morphed faces, they scanned the eyes and upper face more. Local scan structure was not affected by familiarity, but global scan structure was least random (most stereotyped) for novel upright faces. We conclude that expertise (upright faces) leads to less lower-face scanning and more predictable global patterns of information gathering. Experience (famous faces) leads to less upper-face scanning and more idiosyncratic global scan structures, suggesting a superseding influence of facial memories. With morphed faces, subjects return to the upper face to resolve ambiguity, implying a greater importance of this region in face recognition.     

Remembering parts and wholes: Configural processing in face recollection

In two experiments, we examined the role of configural processing on states of awareness in face recognition. Configural processing was manipulated by presenting upright and inverted faces during study and test. After studying facial photographs of humans (Experiment 1) and horses (Experiment 2), participants provided a remember/know judgement for each recognised test face. In both experiments, disrupted configural processing had selective effects on states of awareness, so that inversion reduced remember, but not know, responses. Experiment 2 revealed disproportionate inversion effects in that the difference in remember responses between upright and inverted items was significant for human faces but not horses. These findings suggest that configural processing facilitates explicit recollection by providing a distinctive combination of nonsalient event attributes.     

Effect of inversion on the recognition of external and internal facial features

The purpose of the present study was to find out whether inversion affects recognition of external and internal facial features. 24 participants matched, under two experimental conditions (pair and multiple-choice matchings), upright target faces with three categories of facial test stimuli: full faces, external features and internal features, which were presented in either upright or inverted orientations. Data analysis showed that matching of facial stimuli was faster, more accurate and more consistent under upright than under inverted orientations for all stimulus categories; mostly for full faces, and least for internal features. As a rule, there were no speed-accuracy trade-offs. Implications of the data for accounts of the inversion effect in face recognition in terms of a shift from configurational to componential processing were discussed.     

Orienting to eye gaze and face processing

The author conducted 7 experiments to examine possible interactions between orienting to eye gaze and specific forms of face processing. Participants classified a letter following either an upright or inverted face with averted, uninformative eye gaze. Eye gaze orienting effects were recorded for upright and inverted faces, irrespective of whether the faces were simple, schematic faces or more realistic faces. In contrast, inversion affected orienting to targets appearing along the vertical axis. Switching the contrast between the iris and sclera reversed orienting to eye gaze. Lifting the eyelid to expose more of the iris-sclera contrast led to a potentiation of orienting to eye gaze. Raising the eyebrow alone without the eyelid did not affect orienting. The findings suggest that local perceptual information is critical for orienting to eye gaze and that the effect can occur with a degree of independence from certain types of face processing.     

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.     

An inversion effect modified by expertise in capuchin monkeys

The face inversion effect may be defined as the general impairment in recognition that occurs when faces are rotated 180°. This phenomenon seems particularly strong for faces as opposed to other objects and is often used as a marker of a specialized face-processing mechanism. Four brown capuchin monkeys (Cebus apella) were tested on their ability to discriminate several classes of facial and non-facial stimuli presented in both their upright and inverted orientations in an oddity task. Results revealed significantly better performance on upright than inverted presentations of capuchin and human face stimuli, but not on chimpanzee faces or automobiles. These data support previous studies in humans and other primates suggesting that the inversion effect occurs for stimuli for which subjects have developed an expertise.     

Discrimination of faces and houses by rhesus monkeys: the role of stimulus expertise and rotation angle

The face inversion effect, or impaired recognition of upside down compared to upright faces, is used as a marker for the configural processing of faces in primates. The inversion effect in humans and chimpanzees is strongest for categories of stimuli for which subjects have considerable expertise, primarily conspecifics’ faces. Moreover, discrimination performance decreases linearly as faces are incrementally rotated from upright to inverted. This suggests that rotated faces must be transformed, or normalized back into their most typical viewpoint before configural processing can ensue, and the greater the required normalization, the greater the likelihood of errors resulting. Previous studies in our lab have demonstrated a general face inversion effect in rhesus monkeys that was not influenced by expertise. Therefore, the present study examined the influence of rotation angle on the visual perception of face and nonface stimuli that varied in their level of expertise to further delineate the processes underlying the inversion effect in rhesus monkeys. Five subjects discriminated images in five orientation angles. Results showed significant linear impairments for all stimulus categories, including houses. However, compared to the upright images, only rhesus faces resulted in worse performance at rotation angles greater than 45°, suggesting stronger configural processing for stimuli for which subjects had the greatest expertise.     

Neural developmental changes in processing inverted faces

We explored developmental changes in neural substrates for face processing, using fMRI. Children and adults performed a perceptual-matching task with upright and inverted face and animal stimuli. Behaviorally, inversion disrupted face processing more than animal processing for adults and older children. In line with this behavioral pattern, the left middle occipital gyrus showed a stronger face than animal inversion effect in adults. Moreover, a superior aspect of this region showed a greater face inversion effect in older than in younger children, indicating a developmental change in the processing of inverted faces. The visual regions recruited for inverted face processing in adults also overlapped more with brain regions involved in the viewing of upright objects than with regions involved in the viewing of upright faces in an independent localizer task. Hence, when faces are inverted, adults recruit regions normally engaged for recognizing objects, possibly pointing to a role for the featural processing of inverted faces.