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Reflexive gaze following in common marmoset monkeys

27 Jun 2019-bioRxiv (Cold Spring Harbor Laboratory)-pp 682971

TL;DR: Common marmosets follow the head gaze of conspecifics in order to establish joint attention and this support the assumption of an evolutionary old domain specific faculty shared within the primate order and underline the potential value of marmoset in studies of normal and disturbed joint attention.
Abstract: The ability to extract the direction of the others gaze allows us to shift our attention to an object of interest to the other and to establish joint attention. By mapping ones own expectations, desires and intentions on the object of joint attention, humans develop a Theory of (the others) Mind (TOM), a functional sequence possibly disrupted in autism. Although old world monkeys probably do not possess a TOM, they follow the others gaze and they establish joint attention. Gaze following of both humans and old world monkeys fulfills Fodors criteria of a domain specific function and is orchestrated by very similar cortical architectures, strongly suggesting homology. Also new world monkeys, a primate suborder that split from the old world monkey line about 35 million years ago, have complex social structures. One member of this group, the common marmoset (Callithrix jacchus), has received increasing interest as a potential model in studies of normal and disturbed human social cognition. Marmosets are known to follow human head-gaze. However, the question is if they use gaze following to establish joint attention with conspecifics. Here we show that this is indeed the case. In a free choice task, head-restrained marmosets prefer objects gazed at by a conspecific and, moreover, they exhibit considerably shorter choice reaction times for the same objects. These findings support the assumption of an evolutionary old domain specific faculty shared within the primate order and they underline the potential value of marmosets in studies of normal and disturbed joint attention.nnHIGHLIGHTSO_LICommon marmosets follow the head gaze of conspecifics in order to establish joint attention.nC_LIO_LIBrief exposures to head gaze are sufficient to reallocate an animals attention.nC_LIO_LIThe tendency to follow the others gaze competes with the attractional binding of the conspecifics facenC_LI
Topics: Joint attention (60%), Gaze (57%), Primate (51%)

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Scientific RepoRtS | (2019) 9:15292 | https://doi.org/10.1038/s41598-019-51783-9
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Reexive gaze following in
common marmoset monkeys
Silvia Spadacenta
*
, Peter W. Dicke & Peter Thier
*
The ability to extract the direction of the other’s gaze allows us to shift our attention to an object of
interest to the other and to establish joint attention. By mapping one’s own intentions on the object of
joint attention, humans develop a Theory of (the other’s) Mind (TOM), a functional sequence possibly
disrupted in autism. Gaze following of both humans and old world monkeys is orchestrated by very
similar cortical architectures, strongly suggesting homology. Also new world monkeys, a primate
suborder that split from the old world monkey line about 35 million years ago, have complex social
structures and one member of this group, the common marmosets (Callithrix jacchus) are known to
follow human head-gaze. However, the question is if they use gaze following to establish joint attention
with conspecics. Here we show that this is indeed the case. In a free choice task, head-restrained
marmosets prefer objects gazed at by a conspecic and, moreover, they exhibit considerably shorter
choice reaction times for the same objects. These ndings support the assumption of an evolutionarily
old domain specic faculty shared within the primate order and they underline the potential value of
marmosets in studies of normal and disturbed joint attention.
Gaze following is a an important social skill, which enables humans, great apes and old world primates to establish
joint attention on an object of interest to the other. Although old world monkeys probably do not possess a TOM,
namely they are not able to map ones own intentions, desires and expectations on the object of interest, they fol-
low the others gaze and they establish joint attention
1
. Like humans, gaze following of old world monkeys fullls
Fodors criteria of a domain specic function. Common marmosets are also well known for having a peculiar
interest in faces
2,3
. Unlike macaques, the species of old world primates studied best, and other non-human pri-
mate species, they oen engage in mutual gaze, for example in the context of joint action tasks
4
. Many individuals
even seek eye contact with their human caretakers (personal observations). Common marmosets also care about
the orientation of a human face as demonstrated by the fact that human head-gaze biases choices in an object
selection task
5
. While this latter behavior may indicate an inherent capacity for gaze following, it remains to be
shown that it can also be triggered by conspecics. By the same token the lack of high resolution behavioral data
has as yet precluded well-founded inferences about the relationship of marmoset gaze following to gaze following
exhibited by humans and rhesus monkeys, the two species of old world primates for which detailed behavioral
and neuronal data are available
1,6
. Gaze following of macaques and humans is reex-like in the sense that it is fast
and hard to suppress, two features that have contributed to the assumption of a domain specic faculty
712
based
on a dedicated neural system
13
. Do marmosets follow the gaze of conspecics in the same reex-like manner? An
armative response would support the notion that gaze following in extant primate lines is homologous, i.e. a
reection of shared ancestry.
In order to address these questions, we trained 3 common marmosets (2 females, 1 male) to execute a free
choice task in a well-controlled experimental setup that allowed us to head-restrain the animals to precisely track
eye movements. A conspecic’s face, oriented either to the le or to the right, was presented on a monitor for a
variable time ranging between 100 and 600 ms in steps of 100 ms (Fig.1a) and the observing animal was allowed
to scrutinize the face with eye movements conned by the boundaries of the portrait. e facial portrait was fol-
lowed by the appearance of two targets placed at 5° and + 5° from the center on the horizontal axis. e animals
had to freely choose one of the two targets, a human face (2° × 3°extension), by making an indicative saccade into
a window of 2° centered on the target within 500 ms. Independent of the orientation of the conspecics face, both
possible choices were rewarded, provided that the eyes had met the xation requirements.
Hertie Institute for Clinical Brain Research, Department of Cognitive Neurology, Otfried-Müller-Str. 27, 72076,
Tübingen, Germany. *email: silvia.spadacenta@uni-tuebingen.de; thier@uni-tuebingen.de
open

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Results and Discussion
Common marmosets follow the gaze of a conspecic in a quasi-reexive manner. Figure1b, le
panel, plots the percentage of target choices in the direction of the face orientation (“congruent choices”) as func-
tion of the duration of the availability of the portrait. e graph depicts data pooled over all three animals and the
two possible face orientations: congruent choices exceeded chance level signicantly (binomial probability test,
p < 0.05), indicating that the observing monkey tended to follow the gaze of the portrayed monkey. is prefer-
ence was already apparent aer a presentation duration of the portrait of only 100 ms and got stronger for longer
presentation durations peaking at 300 ms exposure time (see also Supplementary Fig.S1). is dependence on
exposure duration is similar to the one exhibited by human observers when exposed to symbolic central cues
such as pointing arrows. ey typically demonstrate a gradual buildup of their spatial target preferences cued by
central stimuli, reaching an optimum at 300 ms
14,15
. As shown in the right panel of Fig.1b, the dependence of the
choice bias on presentation duration was the same in all three animals for up to 300 ms. Only later, the individual
plots start to diverge: interestingly, two of our animals (M2 and M3) showed a clear second peak, overall convey-
ing the impression of an oscillatory pattern with a period of about 250–300 ms. Periodic uctuations of attention
between two locations with a period of 4 HZ have also been described for human and macaque spatial vision
16,17
.
Figure 1. Oriented faces bias the animal’s choices to targets congruent with gaze direction. (a) Behavioral
paradigm. e trial started with the presentation of a central xation dot. Once xation was established, the
oriented face of a conspecic (replaced by other stimuli in control experiments) appeared for a variable time
(100–600 ms in steps of 100 ms). e disappearance of the conspecic’s face or the control stimuli and the
simultaneous appearance of two peripheral targets was the go signal for the animals to freely choose one of two
peripheral targets presented on the horizontal axis at 5° right and le of the center respectively, by means of a
saccade. e animal received a reward if the xation requirements were met. e xation window for the central
dot had a size of 2° × 2°, a size of 2° × 3° for the peripheral targets portraits of a human and for the central
portraits/control stimuli it corresponded to the extent the centrally presented image (7° × 6°). e subject
depicted in gure has agreed for her portrait to be published. (b) Le panel: plot of the percentage of target
choices congruent with portrait orientation as function of the duration of presentation. Pooled data (monkeys
M1, M2, M3). Binomial probability: ***p < 0.001. Right panel: plot of the percentage of target choices
congruent with portrait orientation for the individual animals M1, M2 and M3. Binomial probability: *p < 0.05,
***p < 0.001. (c) Monkey specic plots of the percentage of target choices congruent with portrait orientation,
separating portraits oriented to the le and right respectively. In each panel (le, M1; center, M2; right, M3) the
solid line stands for le oriented portraits and the dashed one for right oriented portraits.

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Yet, given the fact that the third animal exhibited a dierent pattern, characterized by an absence of a second gaze
following peak and a constant choice at chance level aer 300 ms, further studies will be needed to critically assess
the possibility of periodicity.
All three animals exhibited individually dierent directional biases for the le and the right respectively, mod-
ifying their choice behavior on top of the inuence of directional information provided by facial orientation.
Directional biases became apparent when plotting the dependence of choice preference on head gaze direction
for the three individual animals independently for head gaze to the le and to the right (Fig.1c). For example,
a bias to the le side boosted the correct responses for the le oriented head gaze portraits (M1 and M2, le
and central panel respectively), and for the right oriented when the bias fell on the other side (M3, right panel).
Nonetheless, the bias never altered the overall response curve shape with a peak for congruent choices at around
300 ms. A signicant dominance of congruent choices peaking at 300 ms could be seen in M1 and M2 even for
congruent choices prompted by portraits oriented towards the animals non preferred side (binomial probability,
p < 0.001). A comparable tendency in M3 did not reach signicance (binomial probability, p = 0.1). e basis of
the directional bias remains unclear. e fact that it diers between individuals indicates that hidden imbalances
in the setup that might bind attention can hardly matter.
When the animals were confronted with direct gaze of a conspecic with the face turned straight or alterna-
tively, with black or grey disks of a similar size, likewise lacking directional information, target choices of all three
animals did not dier signicantly from chance level at most exposure times with the exception of the shortest
one (Fig.2a, pooled data). In particular the choice peaks for 300 and 600 ms could no longer been seen. For
100 ms exposure, overall pooled choices to the le were signicantly more frequent than to the right. Two of the
Figure 2. Direct gaze and disk or triangular stimuli attract the animals’ attention towards the center. (a) Plot
of le target choice as function of the duration of a central stimulus, either a conspecics face looking straight
at the observer (direct gaze) or a circular grey or black disk of similar size. Data pooled over the three animals
(monkeys M1, M2, M3). A choice bias is evident at the shortest duration time, whereas for longer exposures
the animals chose the targets on the right and le at random. Binomial probability: ***p < 0.001. (b) Bar plot
of percentage of choices congruent with the darker half of a bipartite disk. Data pooled over the three animals
(monkeys M1, M2, M3). For 100 ms presentation duration, the animals exhibited a signicant preference for the
target on the brighter side, i.e. opposite to the side preference to be expected based on a mechanism exploiting
the luminance asymmetry associated with face orientation. At 300 ms the choices did not indicate a preference.
Binomial probability: **p < 0.01; ***p < 0.001. (c) Bar plot of percentage of choices congruent with the side
cued by the oriented triangles. Data pooled over the three animals (monkeys M1, M2, M3). No signicant bias
was detected.

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individual monkeys (M1, M2) exhibited this preference for the le, whereas the third one (monkey M3) a prefer-
ence for the right. e individual directional preferences for the le and the right corresponded to the direction
of the biases seen in the responses to oriented gaze (Fig.1c), yet, now conned to the shortest exposure only. We
think that the disappearance of the directional bias for longer exposure times might be a consequence of increas-
ing attraction towards the central object, overriding the bias, no matter if the central object is the neutral disk or
the portrait of a conspecic looking straight. is interpretation has interesting implications for the experiments
with oriented faces, which showed a persistence of the directional biases independent of exposure time. Here the
directional gaze seems to suppress the buildup of attraction to the central object, facilitating the readiness to look
elsewhere as determined by the resultant of the others gaze direction and an internal directional bias.
e white ear tus on the le and right of the darker central face of a straight ahead marmoset oer a symmet-
ric luminance prole. Once the animal turns the head to the side, symmetry is lost as the visible area of the ear
tu on the side of the head turn will decrease, whereas the area of the other one will increase (see Supplementary
Fig.S3a,b). Hence, gauging the extent of the luminance asymmetry may be a simple way to determine the others
head gaze direction without the need to process other aspects of the face. To test whether le-right dierences in
the luminance of an object prompt an orienting response of the observer, we exposed all 3 animals to bipartite
disks replacing the marmoset portraits. e disks were black on the le and light grey on the right or vice versa.
ese two versions of the bipartite disks were presented randomly interleaved for 100 ms or 300 ms, two portrait
exposure times that had prompted clear gaze following in the main experiment. Against the backdrop of the
preceding considerations, we had hypothesized that the animals might prefer the target on the side of the darker
half of the bipartite disk for both exposure times. However, contrary to our expectation, the animal preferred the
target on the side of the brighter half of the disk, independently if positioned on the right or le side and, more-
over, only for 100 ms exposure time. For 300 ms choices did not exhibit any preference (Fig.2b). is result does
not support the hypothesis that marmoset gaze following is determined by a simple mechanism, conned to the
comparison of the two ear tu areas.
Turning the head to the side will not only change the balance between paler and darker parts of the face but
also disrupt the symmetry of the facial contour. A face that appears symmetric when seen in a straight ahead
orientation will become asymmetric when turned to the side, an asymmetry that oers directional information.
If marmoset gaze following relied on an ability to exploit contour asymmetries, one might expect that even asym-
metric non-face shapes might redirect attention. In order to test this possibility we exposed our experimental
animals to the vision of lled out black triangles, covering an area similar to the one covered by the portraits. e
triangles were presented with one of their three corners pointing either to the le or to the right. e orientation
was chosen randomly interleaved and the triangles were presented for 100 ms or 300 ms in separate blocks of
trials, i.e. for the presentation durations that had provided the strongest eects of oriented marmoset faces. e
results clearly showed that both for the le and the right oriented triangle the target choice remained at chance
level independent of the presentation duration (see Fig.2c). In sum, the luminance and shape control stimuli,
mimicking isolated low level features of oriented faces, turned out to be unable to redirect spatial attention. In
other words, marmoset gaze following seems to require a more holistic perceptual interpretation of faces, based
on the integrative extraction of more than an isolated parameter.
Congruent choices are accompanied by faster reaction times already at short exposure
times. In the main experiment, the latencies of saccades indicating congruent choices were shorter than the
ones for incongruent choices for exposure times up to 400 ms duration (Fig.3). Actually, this facilitation eect
was strongest for the shortest exposure time, gradually decreased with exposure time and no longer reached sig-
nicance for the longest durations tested (500 and 600 ms; see Fig.3 legend for statistics), consistent with a time
course of reexive rather than volitional orienting. A similar facilitation eect for comparably short exposure
durations has been seen in studies of macaque monkeys
11
and humans
8
. However, these studies did not report a
gradual increase of reaction times with the time of exposure seen in our experiments on marmosets. is dier-
ence might be a consequence of the specic paradigm we used. In our experiments, the animals had to choose
between two targets of equal appearance, rather than to follow the other’s gaze to a specic target as in the work
on macaque monkeys and humans. Hence, our animals may have tended to extract additional information from
the other’s face beyond gaze direction in an attempt to facilitate their choices, provided that this portrait was avail-
able long enough. is increased interest in the other’s face, gated by longer exposure times, can be expected to
compromise the ability to quickly disengage attention at the time of the go-signal. is interpretation is supported
by the experiments with control stimuli and the eye movements prompted by the appearance of the portraits we
discuss below.
Saccades associated with the straight ahead face (“direct gaze”) exhibited latencies that were not dierent from
the ones associated with incongruent choices to oriented faces. Interestingly, latencies of saccades associated with
neutral disks showed an inuence of exposure time that was qualitatively opposite to the inuence on saccades for
congruent choices: while being similar to saccades for straight faces for short presentation durations, they became
shorter with increasing exposure time (see Fig.3). e same held for the bicolor disk control stimuli and for the
oriented triangles (Supplementary Fig.S2). ese results indicate that for marmosets, the attraction of the other’s
face but not the attraction of non-biological objects increases with time exposure and correspondingly attentional
disengagement is delayed. Neutral objects were associated with relatively long saccadic reaction times when pre-
sented briey, probably because of the need to scrutinize the object in order to assess its signicance. Once its
irrelevance is established aer some 200 ms of presentation, the observer disengages her/his attention in order to
prepare a fast saccadic choice. A short exposure to the oriented face can cause a profound shortening of saccadic
reaction time, because the drive to follow gaze direction is already fully expressed whereas facial attraction is still
building up. e idea that the development of facial attraction and in general the perception of faces may need a
much longer time is also supported by a consideration of the pattern of saccadic exploration of the portraits (see

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Supplementary Fig.S3a,b for details), whose complexity keeps growing with exposure time. Hence, the question
is why the drive to follow gaze is fully expressed in saccadic reaction times for short exposure times, arguably
too short to allow a detailed scrutiny of the face whereas the choice bias increases further with exposure time for
up to 300 ms. We think that this dissociation between reaction times and choice probabilities might reect the
concerted action of two systems controlling gaze following. e rst is fast, probably subcortical, controlling gaze
following based on a rough and potentially error prone analysis of the other’s face, too limited to provide informa-
tion on other aspects of the face like the identity or mood of the agent. With longer exposure and concomitantly
processing time, this information becomes available, on the one hand binding attention but, on the other hand,
also improving the directional precision of decisions.
Conclusions
Gaze following is prevalent among numerous species but its strength and exibility varies substantially between
them
18
. As shown here, gaze following is also well developed in common marmosets, a new world monkey spe-
cies. Marmoset gaze following is characterized by strong similarities with the gaze following behavior of the two
old world primate species studied extensively, macaques and humans. e strongest argument for correspond-
ence is the similar dependence on the time of exposure to the others gaze direction. In all three species the other’s
gaze biases decisions on potential targets already aer only 100 ms of exposure to the other’s gaze, too short to
accommodate a more detailed scrutiny of the others face. However, given more time to explore the other’s face,
the bias gets stronger in all three, in line with the assumption that primate gaze following is a faculty, consisting of
an early reex-line component that is complemented by a later, more exible component, arguably also respon-
sible for the more sophisticated emotional and cognitive control known to modulate gaze following
1921
. e
behavioral similarities between the gaze following behavior of marmosets, macaque monkeys and humans are
in principle in line with the assumption of a homologous faculty, already available before the split of the new and
old world monkey primate lines. is conclusion may strengthen the view that the marmoset may indeed become
a useful model system for research into the underpinnings of disturbed human social interactions like autism,
related to decient gaze following and joint attention
22
. However, although compelling, the behavioral similarities
established in our study may as well reect behavioral convergence. Hence, comparative physiological and genetic
studies of the underlying neural systems will be needed to strengthen the case for homology.
Materials and Methods
Experimental model and subject details. Common marmosets. We trained 3 adult common marmoset
monkeys (Callithrix jacchus; two females and one male, aged 7 years) to voluntarily enter a custom made monkey
chair by means of positive reinforcement training and to accept the restriction of head movements through a head
holder. Animals were all born in captivity and kept in a marmoset husbandry at approximately 26 °C, 40%–60%
relative humidity and a 12 h:12 h light-dark cycle. Access to water was always ad libitum, while food intake was
controlled according to body weight (weight loss never less than 10% of the ad libitum weight) and amount of
reward received in the experiment. Food consisted in fresh fruits and vegetables and standard commercial chow.
Figure 3. Oriented faces speed up reaction times for congruent choices. Plots of saccadic reaction times as
function of the duration of presentation of the central stimuli. Data pooled over the three animals (monkeys
M1, M2, M3) and choice direction. Saccadic reaction times indicating congruent choices were signicantly
shorter compared to the incongruent ones up to 400 ms of presentation duration (Wilcoxon rank-sum test,
100 ms: zval = 4.8221, p < 0.001; 200 ms: zval = 3.8449, p < 0.001; 300 ms: zval = 2.0341, p = 0.04; 400 ms:
zval = 2.4745, p = 0.01). e individual plots are tted with 2nd degree polynomial functions in an attempt to
improve the visibility. e two ts that showed a signicant dependence of saccade latency on the presentation
duration of the central stimulus were the one for congruent choices prompted by oriented faces and the one for
neutral disk stimuli. e former exhibited a gradual increase with duration from a substantially shorted reaction
time for a duration of 100 ms (adjusted r
2
= 0.86). e latter, on the other hand, showed a gradual decrease with
duration (adjusted r
2
= 0.89).

Citations
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Dissertation
01 Aug 2018
Abstract: At what point in development does the capacity to reason about what people think emerge? While developmental psychologists have been investigating this question for more than thirty years, the evidence they have gained so far is conflicting. On the one hand, the results of traditional, direct false-beliefs tests, which involve asking participants how a person with a false belief will act, suggest that most children under four years of age are still unaware that beliefs can be false. On the other hand, false-belief tests using indirect measures, such as, for example, looking times or anticipatory looking, suggest that even infants ascribe false beliefs to other people. As many have noted, these results pose a deep developmental puzzle. In this work, I defend the claim that infants can already reason about beliefs. On the one hand, I argue that alternative interpretations of indirect false-belief tests fall short of the mark. On the other, I argue that the fact that young children fail direct false-belief tests can be explained in either of two ways, both of which are compatible with the claim that the capacity to reason about beliefs emerges early on. The first option is to maintain that young children fail because of performance difficulties. This type of position has been defended by other authors, but I argue that the particular proposal I put forward (which I call the processing-time account) offers a better account of the evidence. In contrast, the second option (which I call they hybrid approach) is one that, to the best of my knowledge, no one else has defended so far. This consists in arguing that direct and indirect false-belief tests recruit distinct cognitive systems, each of which can independently sustain the ability to reason about beliefs, but which follow different developmental trajectories. After exploring these two options, I consider which is best supported by the evidence.

3 citations


Cites background from "Reflexive gaze following in common ..."

  • ...• Rachel Razza and Clancy Blair (2009) conducted a longitudinal study of the relationship between mentalising and social competence....

    [...]

  • ...possible, I will be focusing here on their most recent proposal, advanced by Peipei Setoh, Rose Scott and Renée Baillargeon (2016) and known as the Expanded Processing-Demands (EPD for short) account....

    [...]

  • ...The evidence that Westra (2016b) and Carruthers (2016) discuss falls short of establishing their claim – i....

    [...]


References
More filters

01 Jan 1985
Abstract: Abstract We use a new model of metarepresentational development to predict a cognitive deficit which could explain a crucial component of the social impairment in childhood autism. One of the manifestations of a basic metarepresentational capacity is a ‘theory of mind’. We have reason to believe that autistic children lack such a ‘theory’. If this were so, then they would be unable to impute beliefs to others and to predict their behaviour. This hypothesis was tested using Wimmer and Perner's puppet play paradigm. Normal children and those with Down's syndrome were used as controls for a group of autistic children. Even though the mental age of the autistic children was higher than that of the controls, they alone failed to impute beliefs to others. Thus the dysfunction we have postulated and demonstrated is independent of mental retardation and specific to autism.

6,007 citations


Journal ArticleDOI
TL;DR: A new model of metarepresentational development is used to predict a cognitive deficit which could explain a crucial component of the social impairment in childhood autism.
Abstract: We use a new model of metarepresentational development to predict a cognitive deficit which could explain a crucial component of the social impairment in childhood autism. One of the manifestations of a basic metarepresentational capacity is a ‘theory of mind’. We have reason to believe that autistic children lack such a ‘theory’. If this were so, then they would be unable to impute beliefs to others and to predict their behaviour. This hypothesis was tested using Wimmer and Perner's puppet play paradigm. Normal children and those with Down's syndrome were used as controls for a group of autistic children. Even though the mental age of the autistic children was higher than that of the controls, they alone failed to impute beliefs to others. Thus the dysfunction we have postulated and demonstrated is independent of mental retardation and specific to autism.

5,566 citations


Journal ArticleDOI
Hermann J. Müller1, Patrick RabbittInstitutions (1)
TL;DR: Effects of advance cues indicating the probable locations of targets that they had to discriminate and localize support a model for spatial attention with distinct but interacting reflexive and voluntary orienting mechanisms.
Abstract: To study the mechanisms underlying covert orienting of attention in visual space, subjects were given advance cues indicating the probable locations of targets that they had to discriminate and localize. Direct peripheral cues (brightening of one of four boxes in peripheral vision) and symbolic central cues (an arrow at the fixation point indicating a probable peripheral box) were compared. Peripheral and central cues are believed to activate different reflexive and voluntary modes of orienting (Jonides, 1981; Posner, 1980). Experiment 1 showed that the time courses of facilitation and inhibition from peripheral and central cues were characteristic and different. Experiment 2 showed that voluntary orienting in response to symbolic central cues is interrupted by reflexive orienting to random peripheral flashes. Experiment 3 showed that irrelevant peripheral flashes also compete with relevant peripheral cues. The amount of interference varied systematically with the interval between the onset of the relevant cue and of the distracting flash (cue-flash onset asynchrony) and with the cuing condition. Taken together, these effects support a model for spatial attention with distinct but interacting reflexive and voluntary orienting mechanisms.

1,216 citations


Journal ArticleDOI
Abstract: Normal subjects were presented with a simple line drawing of a face looking left, right, or straight ahead. A target letter F or T then appeared to the left or the right of the face. All subjects participated in target detection, localization, and identification response conditions. Although subjects were told that the line drawing’s gaze direction (the cue) did not predict where the target would occur, response time in all three conditions was reliably faster when gaze was toward versus away from the target. This study provides evidence for covert, reflexive orienting to peripheral locations in response to uninformative gaze shifts presented at fixation. The implications for theories of social attention and visual orienting are discussed, and the brain mechanisms that may underlie this phenomenon are considered.

1,094 citations


Journal ArticleDOI
Jon Driver1, Greg Davis, Paola Ricciardelli, Polly Kidd  +2 moreInstitutions (1)
Abstract: This paper seeks to bring together two previously separate research traditions: research on spatial orienting within the visual cueing paradigm and research into social cognition, addressing our tendency to attend in the direction that another person looks. Cueing methodologies from mainstream attention research were adapted to test the automaticity of orienting in the direction of seen gaze. Three studies manipulated the direction of gaze in a computerized face, which appeared centrally in a frontal view during a peripheral letter-discrimination task. Experiments 1 and 2 found faster discrimination of peripheral target letters on the side the computerized face gazed towards, even though the seen gaze did not predict target side, and despite participants being asked to ignore the face. This suggests reflexive covert and/or overt orienting in the direction of seen gaze, arising even when the observer has no motivation to orient in this way. Experiment 3 found faster letter discrimination on the side the computerized face gazed towards even when participants knew that target letters were four times as likely on the opposite side. This suggests that orienting can arise in the direction of seen gaze even when counter to intentions. The experiments illustrate that methods from mainstream attention research can be usefully applied to social cognition, and that studies of spatial attention may profit from considering its social function.

942 citations


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20181