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Abstract Psychological research has long acknowledged that facial expressions can implicitly trigger affective psychophysiological responses. However, whether verbal information can alter the meaning of facial emotions and corresponding response patterns has not been tested. This study examined emotional facial expressions as cues for instructed threat-of-shock or safety, with a focus on defensive responding. In addition, reversal instructions were introduced to test the impact of explicit safety instructions on fear extinction. Forty participants were instructed that they would receive unpleasant electric shocks, for instance, when viewing happy but not angry faces. In a second block, instructions were reversed (e.g., now angry faces cued shock). Happy, neutral, and angry faces were repeatedly presented, and auditory startle probes were delivered in half of the trials. The defensive startle reflex was potentiated for threat compared to safety cues. Importantly, this effect occurred regardless of whether threat was cued by happy or angry expressions. Although the typical pattern of response habituation was observed, defense activation to newly instructed threat cues remained significantly enhanced in the second part of the experiment, and it was more pronounced in more socially anxious participants. Thus, anxious individuals did not exhibit more pronounced defense activation compared to less anxious participants, but their defense activation was more persistent.
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SCIentIFIC RepoRTS | (2018) 8:14988 | DOI:10.1038/s41598-018-33269-2
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Verbal instructions override the
meaning of facial expressions
Florian Bublatzky1,2, Pedro Guerra3 & Georg W. Alpers2
Psychological research has long acknowledged that facial expressions can implicitly trigger aective
psychophysiological responses. However, whether verbal information can alter the meaning of facial
emotions and corresponding response patterns has not been tested. This study examined emotional
facial expressions as cues for instructed threat-of-shock or safety, with a focus on defensive responding.
In addition, reversal instructions were introduced to test the impact of explicit safety instructions on
fear extinction. Forty participants were instructed that they would receive unpleasant electric shocks,
for instance, when viewing happy but not angry faces. In a second block, instructions were reversed
(e.g., now angry faces cued shock). Happy, neutral, and angry faces were repeatedly presented, and
auditory startle probes were delivered in half of the trials. The defensive startle reex was potentiated
for threat compared to safety cues. Importantly, this eect occurred regardless of whether threat
was cued by happy or angry expressions. Although the typical pattern of response habituation was
observed, defense activation to newly instructed threat cues remained signicantly enhanced in the
second part of the experiment, and it was more pronounced in more socially anxious participants.
Thus, anxious individuals did not exhibit more pronounced defense activation compared to less anxious
participants, but their defense activation was more persistent.
e ability to communicate about future events and their potential consequences is highly advantageous for gain-
ing benets and avoiding danger. Such vital information can be transmitted using non-verbal communication
(e.g., facial expressions, body posture)1,2, but also via verbal or written instructions (e.g., ‘beware of ’). Both
sources of information – visual facial expressions and language – have been shown to eectively modulate the
activity of motivational systems in the brain, to prepare for adequate responding in a given situation37. However,
to what degree facial expressions and verbal instructions interact in guiding person perception and social behav-
ior is not well-understood.
ere is a strong body of research examining the role of facial expressions and their capability to mediate
perceptual processing and behavioral responding in social situations. Viewing threat-related emotional expres-
sions – such as fear or anger – has been shown to be associated with enhanced activation of the autonomous
nervous system and speeded behavioral responding810. Similarly, happy facial expressions have been suggested to
receive preferential access to attentional processing resources compared to neutral faces. For instance, happy faces
have been linked to better detection rates11,12 and facilitated electrocortical processing (e.g., LPP component)13.
However, observing unknown people who smile might also be more ambivalent as their actual intention remains
uncertain5,14. Together, these psychophysiological response patterns have been suggested to reect the work-
ings of basic motivational systems that organize behavioral approach or withdrawal (e.g., defense behavior)15,16.
Accordingly, facial expressions of emotion are presumed to be evolutionarily prepared to receive more attentional
resources and prime emotion-specic motor-behavioral responding10,17.
Language is another evolutionary prepared communication system. Aective language, such as insults or
compliments, is especially eective at catching the listeners’ attention. is is particularly evident when infor-
mation directly refers to the listener or reader1820. Accordingly, verbal instructions about imminent aversive
events (threat-of-shock) eectively enhance perceptual processing2123, defensive activation4,7,24,25, and modulate
overt behavioral responding (e.g., in decision-making tasks)26,27. Importantly, this verbal information does not
need to be substantiated by rst-hand experiences of the anticipated aversive events. For instance, despite the
lack of aversive reinforcement, instructed threat contingencies are very resistant to extinction even across several
1Department of Psychosomatic Medicine and Psychotherapy, Central Institute of Mental Health Mannheim, Medical
Faculty Mannheim/Heidelberg University, Mannheim, Germany. 2Clinical Psychology and Biological Psychology
and Psychotherapy, Department of Psychology, School of Social Sciences, University of Mannheim, Mannheim,
Germany. 3University of Granada, Department of Personality, Granada, Spain. Correspondence and requests for
materials should be addressed to F.B. (email: orian.bublatzky@zi-mannheim.de)
Received: 28 February 2018
Accepted: 17 September 2018
Published: xx xx xxxx
OPEN
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SCIentIFIC RepoRTS | (2018) 8:14988 | DOI:10.1038/s41598-018-33269-2
days28. However, verbal instructions can reverse threat expectancies, for instance, when an instructed threat cue
is newly learned as a safety cue2931. Given the centrality of threat perception for interpersonal relations and social
behavior, associations between threatening events and facial information might be malleable and exibly change
according to social settings.
e present study examined the joint impact of visual and verbal aective information on defensive respond-
ing. To this end, pictures of happy and angry facial expressions were verbally instructed as cues for the threat of
electric shocks or safety. As dependent variables, we chose both somatic (startle reex) and autonomic indices
(skin conductance response [SCR] and heart rate [HR]), which have been shown to be sensitive to facial expres-
sions and verbal instructions5,9,24. In addition to physiological measures, we also obtained subjective ratings about
the perceived threat, aective valence, and emotional arousal. Following two previous studies that used pictures of
aective scenes as shock cues (i.e., pleasant and unpleasant IAPS pictures)24,32, threat instructions were predicted
to change the inherent valence of facial expressions. is should be evinced by threat-potentiated startle reex,
enhanced SCRs, and potentiated initial HR deceleration, as well as higher threat ratings for threat-of-shock rela-
tive to safety cues4,7,24,25.
Focusing on the interaction of facial emotions and verbal threat/safety instructions, the congruency of
aective information (e.g., an angry face instructed to signal shock threat compared to safety) was of particular
interest. According to the motivational priming theory15,16, an interaction of threat/safety instruction by facial
expression was expected: When serving as a threat cue, inherently unpleasant stimuli (i.e., angry faces) will elicit
more pronounced defensive responding than inherently pleasant stimuli (i.e., happy faces). Alternatively, incon-
gruent information might be particularly eective in guiding defense activation to unknown people. For instance,
a smiling person instructed to signal shocks may be considered as particularly dangerous14, with implications for
social interactions and behavior towards this person (e.g., impression formation, social bonding)33,34.
Also, we expected to gain insight into the malleability of instruction eects by examining reversal learning.
Reversal learning reects the shi of threat associations from one stimulus to another, with the concurrent inhi-
bition (previous threat cue becomes safe) and acquisition of threat contingencies (a previous safety cue becomes
threatening)35. To this end, a second experimental block was preceded by additional instructions, which aimed
at reversing previously learned threat/safety contingencies (e.g., from threat to safety or vice versa)36,37. Here,
it is of interest whether the impact of reversal instructions on fear extinction learning depends on prepared
learning mechanisms in person perception36. Specically, we examined whether threat eects were more stable
when angry (relative to happy) faces served as reversed safety cues (i.e., previously cueing threat). Moreover, we
predicted that pleasant facial expressions might be less eective as a threat cue37, or that they are more quickly
associated with safety in the reversal test.
Methods
Participants. Sample size was determined using G*Power38, which indicated that N = 40 was required to
detect all relevant physiological eects at a medium eect size (f = 0.25, α error = 0.05, power = 0.8, and assumed
correlation of repeated measures = 0.4). is stop rule for data collection was also in line with previous star-
tle studies using emotional facial expression and threat-of-shock instructions3,24,30. Forty healthy volunteers
(10 males) were recruited from the students of the psychology department at the University of Mannheim.
Participants’ age ranged between 17 and 52 (M = 22.7, SD = 7.1) and the sample was within the normal range of
state and trait anxiety (STAI, M = 35.3 and 35.1, SD = 5.8 and 8.8), social anxiety (SPIN, M = 10.9, SD = 8.2), and
depression (BDI, M = 5.9, SD = 6.5). All participants were informed about the general study procedure before
informed consent was obtained. e ethical review committee of the University of Mannheim approved all uti-
lized procedures and methods. Participants received course credits for their participation.
Stimulus materials and presentation. Face pictures were selected from the Karolinska Directed
Emotional Faces (KDEF39), a well-established stimulus set providing pictures of human facial expressions of
emotion. Sixteen actors (eight females) displaying happy, neutral, and angry facial expressions, were selected
based on visual inspection (i.e., seven raters agreed upon the clarity and recognizability of facial expressions). e
KDEF face identiers were af01, af07, af09, af11, af19, af20, af22, af29, am02, am03, am07, am08, am10, am13,
am14, and am25.
All 48 pictures (1024 × 768 pixels) were presented for 6 s separated by variable inter-trial intervals (ITI) rang-
ing from 10 to 15 s to allow response recovery (see Fig.1). To provoke the defensive startle reex, auditory startle
probes (white noise, 105 dB, 50 ms) were presented during half of the picture trials. e 48 pictures (including the
24 picture-startle trials) were evenly distributed across two experimental blocks (instantiation, reversal) and three
facial expressions (happy, neutral, angry), resulting in four picture-startle trials for each experimental condition
per participant. To prevent the predictability of auditory stimulation, startle probes were presented at either 4,
4.5, 5 or 5.5 s aer picture onset (i.e., while the picture was still visible), and six additional startle probes (three
per block) were presented during the ITI. Startle probes were presented binaurally using headphones (AKG K44
Perception) and the average lag between probes was 28.8 s.
Presentation soware (Neurobehavioral Systems, Inc., Albany, CA, USA) served to control the stimulus pres-
entation, which was pseudorandom regarding picture sequence (no immediate repetition of the same face actor,
no more than three pictures of the same facial expression in a row) and regarding startle presentation (no more
than two picture-startle trials in a row). Electric stimuli for the shock work-up procedure were presented using a
Digitimer Stimulator DS-5 (up to 10 shocks, with maximal 10 mA, 100 ms).
Experimental task and instructions. Participants’ task was to look at all pictures, which were presented
during the two experimental blocks (instantiation and reversal; see Fig.1). Immediately before the rst block
started (instantiation), participants were verbally instructed that they might receive up to three electric shocks
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under specic conditions. One group of participants (N = 20) was told that electric shocks might be administered
whenever an angry face is presented (angry = threat) but not when they see a happy face (happy = safety). e
other group (N = 20) received the opposite instruction, stating that happy facial expressions cued threat-of-shock
(happy = threat), and safety condition being signaled by angry faces (angry = safety). For the second experi-
mental block (reversal), all participants were verbally instructed that now threat and safety contingencies were
reversed. Specically, the previous threat cue becomes safe, and the previous safety cue becomes threatening.
us, across both experimental groups, happy and angry facial expressions served equally oen as instructed and
reversed threat and safety cue; neutral faces always signaled safety.
Procedure. Participants completed questionnaires on general and social anxiety and depression (State-Trait
Anxiety Inventory [STAI-state/trait], Social Phobia Inventory [SPIN], Social Interaction Anxiety Scale [SIAS],
Beck Depression Inventory [BDI]). Sensors for physiological recordings were attached, and an electric stim-
ulation electrode was placed at the right upper arm. Next, a brief shock work-up procedure (without picture
presentation) was carried out to ensure the credibility of the threat instruction22,40. To set the shock intensity
individually at a level rated as “maximally unpleasant but not yet painful”, participants received up to 10 shocks
with increasing intensity. Participants were then instructed that the intensity of the electric shocks given during
the experiment would be equal to the most unpleasant test stimulus.
Practice trials served to familiarize participants with the picture and startle presentation procedure and to
allow for initial habituation of the startle reex. Aerward, verbal instructions regarding threat and safety con-
tingencies were given (i.e., which facial expression signals threat-of-shock and which signals safety) and the
rst experimental block started (instantiation). Following this block, participants rated the hedonic valence and
arousal using the Self-Assessment Manikin (SAM)41, and perceived threat of the facial expressions using a visual
analog scale ranging from not at all to highly threatening (1 to 10). en all participants received the instruction
that threat/safety contingencies were now reversed (e.g., the threat cue becomes safe, and safety cue becomes
threatening), and the second block started (reversal). Facial expressions were rated again aer the reversal block.
Finally, participants were debriefed. No shocks were presented during the experiment. us, results reect phys-
iological responding during the anticipation (but not experience) of electric shocks.
Data recording and reduction. Psychophysiological measures were recorded continuously with a vAmp
amplier (BrainProducts, Munich, Germany). Startle amplitudes were derived from the electromyogram of the
orbicularis muscle using two miniature Ag/AgCl electrodes. e raw signal was recorded at a 1000 Hz sampling
rate and frequencies below 28 Hz and above 500 Hz were ltered out with a band-pass lter (24 dB/octave roll-o).
Raw electromyogram (EMG) data were rectied and smoothed with a moving average procedure (50 ms) in
VisionAnalyzer 2.0 (BrainProducts). Startle responses were scored with an automated procedure and dened as
the maximum peak in the 21–150 ms time window following each startle probe. Peak amplitudes were calculated
relative to a mean baseline period (50 ms preceding startle response time window)28,42.
Figure 1. Schematic illustration of the experimental procedures and stimulus presentation. (A) Aer a
brief practice run and shock work-up procedure, participants were verbally instructed that one particular
emotional facial expression serves as a cue for threat-of-shock (e.g., happy) or safety (e.g., angry) and the rst
experimental block started (instantiation). Preceding the second experimental block (reversal), a verbal reversal
instruction stated that now threat and safety contingencies are reversed (e.g., now angry faces cue threat and
happy cue safety). e order in which facial expressions cued threat or safety was tested in two groups (each
N = 20 completed the happy-angry or angry-happy threat order). Please note, neutral faces always cued safety.
Following each block, threat and safety cues were rated regarding valence, arousal, and perceived threat. (B)
Within each block, face pictures displaying happy, neutral, and angry facial expressions were presented (each
6 s) with a variable intertrial interval (ITI, 10 to 15 s). Auditory startle probes were presented occasionally
during pictures and ITIs, no shocks were presented during the experiment. Example pictures are taken from the
KDEF (identiers: af01has, am08ans, am10nes, and af20ans).
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As an index of phasic autonomic activation, skin conductance responses (SCRs) were recorded with Ag/
AgCl electrodes (constant voltage of 0.5 V; 20 Hz sampling rate) placed at the hypothenar eminence of the
non-dominant hand. SCRs to picture onset were calculated as the maximum increase in skin conductance in
the interval of 1 to 6 s (relative to a 1 s pre-stimulus period). A minimum threshold of 0.02 µS was used for
zero-response detection, and range and distribution corrections were applied. Phasic heart rate changes to picture
onset was derived from the electrocardiogram recorded at lead II. e electrocardiogram signal was recorded at
1000 Hz, and frequencies below 0.1 and above 13 Hz were ltered. e weighted HR averages every half second
were expressed in terms of dierential scores with respect to a 2 s baseline period24.
Data analysis and statistical design. Self-report (valence, arousal, and threat ratings) and physio-
logical data (startle-EMG and SCR) were submitted to (2 × 2) × 2 repeated measures ANOVA, including the
within-subject factors Instruction (threat vs. safety) and Block (instantiation Block 1 vs. reversal Block 2), as well
as the between-group factor Order (happy-angry vs. angry-happy). e Order referred to the sequence in which
emotional facial expression cued threat or safety in which experimental block. Specically, for the happy-angry
order, happy faces served as threat cue during instantiation block (Block 1), and angry faces cued threat during
the following reversal block (Block 2). is was reversed for the angry-happy order, in which angry faces during
Block 1 and happy faces in Block 2 cued threat-of-shock. Regarding phasic changes in heart rate, an additional
factor, Time (12), was implemented to compare half-second changes aer picture onset.
To examine the impact of emotional facial expression on the instantiation and reversal of threat instruc-
tions, planned comparisons focused separately on each Order (happy-angry vs. angry-happy). Please note that
for reasons of brevity and to reduce the complexity of the overall design, neutral faces cued safety in both blocks
and were thus excluded from the analyses of instructed and reversed threat. However, supplementary analyses
were conducted to compare Facial expression (happy vs. neutral vs. angry) when serving as a safety cue (see
Supplemental Material). Covariation analyses were conducted to test the impact of inter-individual dierences in
reported social- and trait-anxiety on defense activation.
Greenhouse-Geisser corrections were used where relevant, and the partial ƞ2 is reported as a measure of eect
size. To control for Type 1 error, Bonferroni correction was applied for post-hoc t-tests.
Results
Self-report data. Threat ratings. Overall, instructed threat cues were rated as more threatening rel-
ative to safety cues, F(1,39) = 17.37, p < 0.001, ηp2 = 0.31, and perceived threat decreased from the instan-
tiation block to the reversal block, F(1,39) = 5.98, p < 0.05, ηp2 = 0.13. The interaction Instruction × Block
did not reach significance, F(1,39) = 1.47, p = 0.23, ηp2 = 0.04, however, a significant three-way interaction
Instruction × Block × Order emerged, F(1,38) = 56.72, p < 0.001, ηp2 = 0.60.
Follow-up analyses run separately for the Happy-Angry order (see Fig.2A; Table1) showed that instructed
threat eects varied across blocks, F(1,19) = 21.26, p < 0.001, ηp2 = 0.53. Specically, pronounced threat rat-
ings were observed for happy facial expressions cueing threat-of-shock during instantiation, F(1,19) = 5.23,
p < 0.05, ηp2 = 0.22, and for angry faces cueing threat in the subsequent reversal block, F(1,19) = 31.30, p < 0.001,
ηp2 = 0.62. Similarly, for the Angry-Happy order, an interaction Instruction × Block emerged F(1,19) = 35.46,
p < 0.001, ηp2 = 0.65. Separate comparisons of threat and safety cues indicate more pronounced threat ratings for
angry faces during the instantiation block, F(1,19) = 35.04, p < 0.001, ηp2 = 0.65, and happy expressions in the
reversal block, F(1,19) = 16.50, p < 0.01, ηp2 = 0.47.
Valence ratings. Overall, threat cues were rated as more unpleasant than safety cues, F(1,39) = 11.01, p < 0.01,
ηp2 = 0.22, and unpleasantness decreased across blocks, F(1,39) = 4.62, p < 0.05, ηp2 = 0.11. Whereas no interac-
tion of Instruction × Block was observed, F(1,39) = 1.99, p = 0.17, ηp2 = 0.05, a signicant three-way interaction
was found, Instruction × Block × Order F(1,38) = 60. 87, p < 0.001, ηp2 = 0.62.
Separate analysis for the Happy-Angry order (Fig.2B) showed a signicant interaction Instruction × Block ,
F(1,19) = 13.53, p < 0.01, ηp2 = 0.42. Interestingly, happy expressions were rated as more pleasant than angry faces
even when happy faces served as instructed threat cue in the instantiation block, F(1,19) = 11.01, p < 0.01, ηp2 = 0.37.
In the reversal block, angry faces cueing threat were rated as more unpleasant than happy faces cueing safety,
F(1,19) = 11.56, p < 0.01, ηp2 = 0.38. For the Angry-Happy order, a signicant interaction Instruction × Block was
evident, F(1,19) = 58.77, p < 0.001, ηp2 = 0.76. Angry faces cueing threat were more unpleasant during the instan-
tiation block, F(1,19) = 72.96, p < 0.001, ηp2 = 0.79, and this threat eect was less pronounced when happy facial
expressions served as new threat cues in the reversal block, F(1,19) = 11.28, p < 0.01, ηp2 = 0.37.
Arousal ratings. Overall, instructed threat cues were rated as more arousing than safety cues, F(1,39) = 10.41,
p < 0.01, ηp2 = 0.21, and arousal decreased from the instantiation to the reversal block, F(1,39) = 21.90, p < 0.001,
ηp2 = 0.36. Moreover, the interaction of Instruction × Block was signicant, F(1,39) = 4.83, p < 0.05, ηp2 = 0.11,
showing pronounced threat eects in the instantiation block, F(1,39) = 13.52, p < 0.01, ηp2 = 0.26, but not in the
reversal block, F(1,39) = 1.05, p = 0.31, ηp2 = 0.03. is pattern did not signicantly dier between experimental
orders (Angry-Happy or Happy-Angry; Fig.2C), Instruction × Block × Order F(1,38) = 2.91, p = 0.10, ηp2 = 0.07.
Startle reex. e defensive startle reex was more pronounced when viewing threat as compared to safety
cues (Fig.3A), F(1,39) = 56.87, p < 0.001, ηp2 = 0.59, and a pronounced pattern of response habituation was
observed across experimental blocks (Fig.4A), F(1,39) = 58.19, p < 0.001, ηp2 = 0.60. Moreover, reex amplitudes
varied as a function of Instruction × Block, F(1,39) = 4.26, p < 0.05, ηp2 = 0.10, indicating that threat-potentiation
decreased from the instantiation to the reversal block. Post-hoc tests revealed pronounced dierences between
threat and safety cues within the instantiation block, and less markedly but still highly signicant in the following
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SCIentIFIC RepoRTS | (2018) 8:14988 | DOI:10.1038/s41598-018-33269-2
reversal block, Fs(1,39) = 48.78 and 16.52, ps < 0.001, ηp2 = 0.56 and 0.30. Importantly, the inherent valence of
emotional facial expressions did not modulate the instantiation and reversal of threat as observed for the startle
reex, Order × Instruction × Block, F(1,38) = 0.73, p = 0.40, ηp2 = 0.02. Planned follow-up tests focused on each
experimental order separately.
Figure 2. Self-reported threat (A), valence (B), and arousal (C) ratings as a function of facial expression (happy,
angry) and instructions (threat, safety). e le side illustrates overall means (with SEM) averaged across
experimental blocks and orders. On the right side, separate means are plotted for each order. e angry-happy
order started with angry facial expression cueing threat during the instantiation block and happy faces cueing
threat during the following reversal block. For the happy-angry order, instructed threat/safety contingencies
were vice versa.
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When angry faces signaled threat during the instantiation block and served as safety cue in the subsequent
reversal block (Angry-Happy order), main eects of Instruction and Block were signicant, Fs(1,19) = 23.67
and 49.31, ps < 0.001, ηp2 = 0.56 and 0.72. Moreover, startle amplitudes tended to vary as a function of
Instruction × Block, F(1,19) = 3.38, p = 0.08, ηp2 = 0.15. Separate analyses for each block revealed that angry faces
as threat cue triggered highly signicant threat eects during instantiation, F(1,19) = 24.66, p < 0.001, ηp2 = 0.57,
but only marginal threat effects were observed when angry faces served as safety cue in the reversal block,
F(1,19) = 4.02, p = 0.06, ηp2 = 0.17. In contrast, for the Happy-Angry order, when happy faces served as threat
cue during instantiation and as safety cue in the reversal block, main eects of Instruction and Block were found,
Fs(1,19) = 19.97 and 32.59, ps < 0.001, ηp2 = 0.51 and 0.63. However, threat eects were not reduced across blocks
when angry faces cued shock threat in the reversal block, Instruction × Blo ck F(1,19) = 0.96, p = 0.34, ηp2 = 0.05.
Follow-up tests revealed pronounced threat-potentiated startle for happy faces cueing threat in the instantiation
block, F(1,19) = 22.95, p < 0.001, ηp2 = 0.55, which was similarly pronounced in the subsequent reversal block
when angry faces cued threat, F(1,19) = 16.1, p = 0.001, ηp2 = 0.46. us, during the reversal block, instruction
eects were more resistant to extinction when angry rather than happy faces cued threat.
Exploratory analyses revealed that the overall interaction Instruction × Block varied as a function of
inter-individual dierences in reported social- and trait-anxiety (Fig.4B). Specically, signicant covariation
eects were observed with SPIN scores, F(1,38) = 8.15, p < 0.01, ηp2 = 0.18, STAI-trait, F(1,38) = 7.81, p < 0.01,
ηp2 = 0.17, and marginally with SIAS, F(1,38) = 3.71, p = 0.06, ηp2 = 0.09. To follow up on these interactions,
correlational analyses were conducted between anxiety scores and startle amplitudes (i.e., the dierence scores
between threat minus safety) separately for each block. For the instantiation block, threat eects did not vary
with anxiety level (rtrait-anxiety = 0.20, p = 0.21; rSPIN = 0.25, p = 0.12; rSIAS = 0.08, p = 0.61). In the subsequent
reversal block, however, threat-potentiated startle was more pronounced in participants who scored higher on
anxiety (rtrait-anxiety = 0.36, p < 0.05; rSPIN = 0.32, p < 0.05; rSIAS = 0.32, p < 0.05). us, anxious participants did not
exhibit more pronounced, but more persistent defense activation compared to less socially anxious participants.
Skin conductance responses. Enhanced skin conductance responses (SCR) were observed for threat rel-
ative to safety cues, F(1,39) = 9.29, p < 0.01, ηp2 = 0.19 (see Fig.3B). SCRs diminished over time, they were more
pronounced in the instantiation block than in the subsequent reversal block, F(1,39) = 7.21, p < 0.05, ηp2 = 0.16.
e interaction Instruction × Block didn’t reach signicance, F(1,39) = 2.95, p = 0.09, ηp2 = 0.07. Explorator y
follow-up analyses revealed signicant threat eects during instantiation, F(1,39) = 20.93, p < 0.001, ηp2 = 0.35,
but not in the reversal block, F(1,39) = 2.31, p = 0.14, ηp2 = 0.06. Importantly, the inherent facial valence did
not modulate SCRs for instantiation and reversal of threat, Order × Instruct ion × Block, F(1,38) = 0.53, p = 0.47,
ηp2 = 0.01. Planned comparisons focused separately on each experimental order.
When angry faces served initially as threat cues, and later as safety cues (Angry-Happy order), there
were signicant main eects of Instruction, F(1,19) = 13.34, p < 0.01, ηp2 = 0.41, and Block, F(1,19) = 8.43,
p < 0.01, ηp2 = 0.31, as well as a trend to an interaction Instruction × Block, F(1,19) = 3.55, p = 0.08, ηp2 = 0.16.
Follow-up analyses revealed threat-enhanced SCRs when angry faces cued threat during the instantiation block,
F(1,19) = 15.15, p = 0.001, ηp2 = 0.44, but not when happy faces cued threat in the reversal block, F(1,19) = 2.63,
p = 0.12, ηp2 = 0.12. For the Happy-Angry order, in contrast, SCRs did not differ for Instruction or Block,
Fs(1,19) = 2.55 and 0.53, ps = 0.13 and 0.48, ηp2 = 0.21 and 0.03. Whereas no interaction of Instruction × Block
was found, F(1,19) = 0.41, p = 0.53, ηp2 = 0.02, exploratory analyses indicated threat-enhanced SCRs to happy
faces cueing threat during the instantiation, F(1,19) = 6.85, p < 0.05, ηp2 = 0.27, but not when threat was cued
by angry faces in the subsequent reversal block, F(1,19) = 0.79, p = 0.38, ηp2 = 0.04. No covariation eects were
observed between SCRs and anxiety scores.
Phasic heart rate changes. Overall, heart rate revealed a signicant deceleration when viewing threat
relative to safety cues, F(1,39) = 4.03, p = 0.05, ηp2 = 0.09 (see Fig.3C). Furthermore, there was an interaction of
Time × Instruction, F(11,429) = 5.48, p < 0.01, ηp2 = 0.12. Follow-up analyses were calculated separately for each
time interval and indicated signicant heart rate deceleration for threat relative to safety cues between 3 and 5 s
Block Instruction Order
Startle SCR HR Val e n ce Arousal reat
MSD MSD MSD MSD MSD MSD
Instantiation
(Block 1)
reat Angry-Happy 57.85 5.14 0.115 0.11 2.75 1.66 3.55 1.73 5.15 1.66 5.05 2.61
Happy-Angry 57.48 5.31 0.097 0.17 2.75 2.71 5.05 2.48 5.80 2.14 4.05 3.61
Safe Angry-Happy 49.54 3.31 0.030 0.04 0.78 1.85 7.90 1.25 3.30 2.13 0.95 1.76
Happy-Angry 49.77 3.60 0.032 0.07 1.30 4.36 3.50 1.67 5.45 2.19 5.20 3.08
Reversal
(Block 2)
reat Angry-Happy 50.26 5.99 0.048 0.10 1.64 3.28 6.30 1.75 4.15 2.08 2.15 1.95
Happy-Angry 51.21 5.44 0.079 0.24 2.58 2.65 4.85 1.69 3.90 1.71 4.10 2.38
Safe Angry-Happy 46.58 4.22 0.014 0.02 1.33 2.62 4.50 1.54 3.75 1.62 4.23 2.46
Happy-Angry 45.41 4.32 0.034 0.05 1.29 2.31 6.70 1.75 3.75 1.89 1.55 1.79
Table 1. Mean amplitudes and standard deviations (M, SD) as a function of Block (instantiation vs. reversal),
Instruction (threat vs. safety), and Order of threat instruction (Angry-Happy vs. Happy-Angry). Means are
provided for the startle reex, skin conductance responses (SCR), heart rate (HR), and ratings of the self-
reported valence, arousal, and perceived threat. Heart rate changes refer to averages across the signicant time
intervals from 3 to 5 s aer picture onset.
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aer picture onset (all ps < 0.05). Neither the main eect Block, F(1,39) < 0.01, p = 0.98, ηp2 < 0.01, nor the inter-
actions Instruction × Block, F(1,39) = 0.78, p = 0.38, ηp2 = 0.02, Time × Instruction × Block, F(11,429) = 1.04,
p = 0.38, ηp2 = 0.03, nor the four-way interaction by Order reached signicance, F(11,418) = 0.65, p = 0.58,
ηp2 = 0.02.
Exploratory analyses focused separately on the dierent experimental orders. For the Angry-Happy order,
when angry faces served as a threat cue in the instantiation block, a substantial threat deceleration was evident,
F(1,19) = 8.98, p < 0.01, ηp2 = 0.32, which developed over time following the threat cue onset, F(11,209) = 5.22,
p < 0.01, ηp2 = 0.22. However, no threat eects emerged in the subsequent reversal block when happy faces served
as new threat cue, F(1,19) = 0.01, p = 0.93, ηp2 < 0.01. For the Happy-Angry order, in contrast, no threat eect
occurred, F(1,19) = 2.62, p = 0.12, ηp2 = 0.12, nor did any interaction including Instruction reach signicance,
Fs < 1.68, ps > 0.20, ηp2 < 0.08. No covariation eects were found between phasic heart rate changes and anxiety
scores.
Figure 3. Mean responses of the startle reex (A) and skin conductance (B) for happy and angry facial
expressions serving as threat or safety cue (with SEM). Heart rate changes (C) are averaged every half a second
aer stimulus onset. As no interaction eects occurred with the sequence of instructions, averages across
experimental orders (happy-angry and angry-happy) are illustrated.
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Discussion
The present study examined the capability of emotional facial expressions as cues for verbally instructed
threat-of-shock or safety. Also, we tested the exibility of threat and safety associations using reversal instructions.
Verbal communication about threat contingencies triggered, as expected, a pronounced pattern of psychophys-
iological defense reactions. is was evident in potentiated eye-blink startle reex, enhanced skin conductance
responses, and heart rate deceleration. For self-report data, interaction eects of facial expressions and verbal
instructions emerged. Specically, when smiling faces cued threat, they were rated as aversive as angry faces.
In contrast, physiological responding was independent of whether the threat was cued by a happy or an angry
facial expression. Moreover, reversal instructions exibly changed defense activation, leading to relatively stable
threat eects despite substantial response habituation across the experimental blocks. Interestingly, aer reversal
instructions, threat-potentiated startle was more pronounced in more socially anxious participants. us, anxious
individuals did not exhibit more pronounced defense activation compared to less anxious participants, but their
defense activation was more persistent.
When confronting other people’s facial expressions, which were previously learned as signals for shock threat,
pronounced activation of the somatic and autonomic nervous system was observed (i.e., potentiated eye-blink
startle and enhanced skin conductance responses). ese ndings replicate previous studies showing defense
activation to visual stimuli cueing instructed threat-of-shock2429. Defensive responding, however, occurred
regardless of whether the threat was cued by a smiling or an angry face. us, the inherent valence of the threat
cue (happy or angry expression) was not relevant for the acquisition of threat contingencies. is nding adds
to previous research using the threat-of-shock paradigm with complex natural aective scenes4,7,24. For example,
Bradley and colleagues24 observed comparable threat-potentiated startle reex to pleasant and unpleasant pictures
when these served as instructed threat cues. Moreover, when pictures were not predictive for threat-of-shock (i.e.,
presented within a threatening context), threat eects were found similarly pronounced for pleasant, neutral,
and unpleasant pictures4,43. e present study extends this view to face and person perception and shows that the
emotional salience of happy and angry facial expressions can be easily overridden by verbal instructions about
threat contingencies. is nding contributes to the rather mixed evidence on whether human faces serve as an
evolutionary prepared conditional stimulus36,37,44,45. Compared to pictures of snakes or spiders, the human face
may be a less reliable source of threat or safety information, probably because facial expressions can be manipu-
lated consciously and are subject to social display rules46.
e inherent valence of an emotional face did not interact with the verbally transmitted acquisition of threat
or safety contingencies. is nding is supported by previous neuroimaging research, for instance, showing
that threat instructions led to a more general sensitization of stimulus processing32,47, regardless of the a priori
Figure 4. (A) Mean startle responses as a function of threat/safety instructions averaged across experimental
blocks and orders (with SEM). reat and safety contingencies were instantiated in Block 1 (e.g., angry faces
cue threat) and reversed in Block 2 (e.g., happy faces cue threat; or vice versa). (B) Scatterplots illustrate the
covariation between individuals’ social anxiety and threat-potentiated startle eects (dierences between threat
and safety) separately for the instantiation and reversal block. Overall, threat eects are malleable and stable;
anxious participants reveal a more persistent pattern of defense activation aer reversal instructions.
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SCIentIFIC RepoRTS | (2018) 8:14988 | DOI:10.1038/s41598-018-33269-2
meaning of a shock cue (e.g., unpleasant or neutral social scenes). Moreover, in the present study, neither the
somatic (eye-blink startle) nor the autonomic nervous system (SCR and phasic HR responses) showed an interac-
tion between visual and instructed information. Supplementary analyses using Bayesian statistics supported these
ndings. Likelihood estimates of the null hypothesis (i.e., no Order × Instruction × Block interaction) indicated
that the null relative to the alternative hypotheses were around 19-, 37-, and 142-times more likely for the startle
reex, SCR, and HR measures respectively. us, the present data lend support for the notion that the processing
of visual and verbal threat information is organized in (partially) distinct neural circuitry. For example, aective
modulation of the startle reex triggered by emotional pictures is impaired in patients with right rather than le
temporal lobectomy, whereas the opposite pattern can be observed when instructed threat cues are presented48.
Interestingly, our self-report data revealed result patterns that were partly in contrast to physiological measures.
Valence, arousal, and threat ratings conrmed that verbal communication about potential threats clearly induced
aversive anticipations. Moreover, these threat/safety contingencies were highly malleable and reversible using
subsequent instructions. In contrast to physiology, however, rating data showed that the impact of threat and
safety instructions varied with the inherent valence of the facial threat/safety cue. When cueing threat, a smiling
face becomes as aversive as an angry face, and both cues were highly eective in triggering defensive responding
to cope with the anticipated aversive event.
Overall, reversal instructions exibly changed threat/safety associations and the corresponding physiological
response patterns. In line with previous studies, verbal instructions were highly eective at reducing defensive
responding using reversing aective contingencies from threat to safety2931. Similarly eective was the rever-
sal of contingencies from safety to threat. Newly learned threat cues (previously safe), compared to the newly
learned safety cues (previously threatening), were associated with lower valence and higher threat ratings.
Moreover, potentiation of the startle reex was observed for the new threat cues despite pronounced response
habituation across experimental blocks. is result adds to the ndings of previous research, which show that
instructed threat eects may be highly persistent within and across repeated sessions, even without any aversive
reinforcement4,28.
Interestingly, after reversal instructions, threat effects varied as a function of social and trait anxiety.
Specically, anxious participants did not exhibit more pronounced defense activation compared to less socially
anxious participants but did exhibit a more persistent defense activation. From a clinical perspective, this is an
important nding showing that inter-individual dierences in anxiety might account for the capability to learn
new safety contingencies and to reduce psychophysiological defense activation. As many fears and anxieties rely
on aversive anticipations rather than experiences, the mere absence of aversive events or omission of reinforce-
ment is not sucient for successful fear extinction learning (e.g., in generalized anxiety disorder or social pho-
bia)4951. To optimize social communication about threats and safety in a therapeutic context, dierent means
of social learning need to be accounted for (i.e., learning by instructions and observations)25,52. Building upon
the present inter-individual dierences in reversal learning, testing (sub-)clinical samples high in social anxiety
or interpersonal disturbances might be particularly informative53,54. Here, the implementation of a full reversal
design29,35 might focus on safety learning and elucidate the impact of reversed compared to maintained social
safety cues.
Several noteworthy aspects of the present design and findings need to be acknowledged and should be
addressed in future research. Exploratory analyses provided some indication for the hypothesis that facial
emotions might dierentially modulate reversal learning. Specically, for the startle reex during the reversal
block, instruction eects were more resistant to extinction when angry rather than happy faces cued threat. is
nding might result from anger-superiority in threat learning (i.e., angry faces more readily associated with
threat)9,10 or happy-superiority11,12 in safety learning. For directly comparing these opposing hypotheses, the use
of a non-aective threat cue condition would have been useful (i.e., neutral faces cueing threat during reversal
block) and cannot be resolved with the data at hand. Future research could examine the capability of distinct
non-aective social stimuli as reversed threat/safety cue. For instance, invariant facial features – such as person
identity and the color of the skin – have been shown to be powerful factors that bias threat learning and can be
pitted against each other (e.g., viewing other-race, but same team faces)34,36,45. Here, social approaches to initiate
persistent reversal learning (i.e., shiing aversive contingencies to other non-social cues) may help to counteract
stereotypes, social avoidance, and ostracism35,55. From an evolutionary perspective, it appears likely that com-
bined variant and invariant facial information (e.g., facial expression and person identity cues)56 critically guide
behavioral responding. For instance, an angry looking out-group member or a smiling mother might be more
readily learned as a signal for threat or safety; such congruency eects in prepared learning can be tested with
personalized stimulus materials (e.g., pictures of attachment gures)57,58. Finally, the transfer to behavioral output
measures appears pertinent to test the implications and consequences of threat and safety learning in social inter-
action situations, for instance, regarding interpersonal trust59, stereotyping and social group biases33,34, or choice
behavior in clinical settings (e.g., decision to undergo treatment)54,60.
In summary, verbal communication about threats might easily prime defensive response programs regardless
of the inherent valence of the threat or safety cue (i.e., happy or angry facial expression). Moreover, threat eects
were malleable by additional verbal instructions, and the persistence of threat eects varied with inter-individual
dierences in social and trait anxiety. Anxious participants did not exhibit more pronounced defense activation
compared to less anxious participants but did exhibit more persistent defense activation. As threat instructions
were not substantiated by the individual’s own experiences (i.e., no shocks during the experiment), these ndings
demonstrate the eects of mere anticipatory processes in person perception relevant to maladaptive extinction
learning in anxiety disorders.
Data Availability
e datasets generated and analyzed during the current study are available from F.B on request.
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10
SCIentIFIC RepoRTS | (2018) 8:14988 | DOI:10.1038/s41598-018-33269-2
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Acknowledgements
We are grateful to O. Denzler, O. Grüner, S. Reuter, and C. Zala for their help with data collection. We received
helpful comments from Fatih Kavcioglu. is research was supported by the German Research Foundation
(Deutsche Forschungsgemeinscha; BU 3255/1-1, granted to F. Bublatzky).
Author Contributions
F.B., P.G. and G.W.A conceived the study and were involved in the generation and analyses of the data. F.B. wrote
the manuscript and all authors revised the manuscript.
Additional Information
Supplementary information accompanies this paper at https://doi.org/10.1038/s41598-018-33269-2.
Competing Interests: e authors declare no competing interests.
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Supplementary resource (1)

... Forty-five students (36 female, mean age = 20.04 years, SD = 1.93) were recruited from the University of Granada (Spain). Sample size was chosen similar to previous research using facial expressions and instructed threat manipulations 11,12,28,33,34 and is in line with estimations based on G*power 35 . Statistical estimations indicate that N = 46 is required to detect instruction by face category interaction effects at a medium effect size (f = 0.20, power = 0.90, α error = 0.05, and assumed correlation of repeated measures = 0.5). ...
... For instance, Costa et al. 27 showed that neutral stimuli associated with threat-of-shock or safety can be reversed from cueing threat to safety and vice versa. Similarly, verbal threat instantiation and reversal instructions can readily override the implicit affective meaning of emotional facial expressions (e.g. a smile may also signal threat 28,33 ). Importantly, however, reversal learning implicates the workings of (at least) two concurrent processes: the inhibition of previously learned threat-associations, while a new threat-association is established 51 . ...
... Whereas divergent findings may relate to different tasks (passive viewing vs. instructed threat) and/or reduced trial numbers, other alternative hypotheses are of interest. Specifically, an over-generalization of threat might have occurred across face categories 62 , and/or overwritten the implicit affective picture qualities through verbal instructions 28,33 . This also relates to clinical phenomena, which are observable, for example, in the emergence and treatment of phobias, panic, or trauma-related disorders. ...
Article
Full-text available
Significant others provide individuals with a sense of safety and security. However, the mechanisms that underlie attachment-induced safety are hardly understood. Recent research has shown beneficial effects when viewing pictures of the romantic partner, leading to reduced pain experience and defensive responding. Building upon this, we examined the inhibitory capacity of loved face pictures on fear learning in an instructed threat paradigm. Pictures of loved familiar or unknown individuals served as signals for either threat of electric shocks or safety, while a broad set of psychophysiological measures was recorded. We assumed that a long-term learning history of beneficial relations interferes with social threat learning. Nevertheless, results yielded a typical pattern of physiological defense activation towards threat cues, regardless of whether threat was signaled by an unknown or a loved face. These findings call into question the notion that pictures of loved individuals are shielded against becoming threat cues, with implications for attachment and trauma research.
... flexibly changed by means of new information (e.g., reversal instructions; Atlas, 2019;Bublatzky et al., 2018Bublatzky et al., , 2019. Focusing on the interaction of face identity and facial emotion, the present study examined face and person perception as a function of verbal learning. ...
... Such attentional threat biases provide the perceptual base for organizing adaptive behaviors towards signals of threat and harm. For instance, verbally instructed threat linked to facial stimuli has been shown effective to provoke enhanced activity of the somatic and autonomic nervous system (e.g., potentiated startle reflex and enhanced skin conductance responses; Grillon & Charney, 2011;Bublatzky et al., 2018Bublatzky et al., , 2019. This physiological priming of defensive response systems is further associated with overt avoidance behaviors and anxious decisions for costly but safe behavioral choices Pittig, Treanor et al., 2018;Pittig, Hengen et al., 2018). ...
... As in previous research, our rating data demonstrate that instructions readily instantiated and reversed threat/safetyassociations. On the other hand, reversal instructions have been shown to effectively modulate the activity of the autonomic and somatic nervous system. For instance, threatenhanced skin conductance responses and startle reflex activity indicated reversed fear responses (Atlas & Phelps, 2018;Bublatzky et al., 2018Bublatzky et al., , 2019Costa et al., 2015;Mertens et al., 2018). Extending these findings, the present study revealed reversal-related brain activity specifically to newly learned threat and safe identities over visual processing areas at around 200 msec after the onset of a face picture (N170 and EPN). ...
Article
The face of a friend indicates safety, the face of a foe can indicate threat. Here, we examine the effects of verbal instructions (‘beware of this person’) on the perception of unknown persons. Focusing on visual attention, face identity and facial expression information is examined during instructed threat-of-shock or safety. However, shocks never occurred. Participants quickly acquired instructed threat associations, and electrocortical processing differentiated threat-from safe-identities as well as emotional and neutral facial expressions. Importantly, face encoding varied as a joint function of identity and facial expression, as revealed by pronounced N170 amplitudes to smiling threat-identities. Moreover, instructions readily reversed previously learned affective associations leading to attention allocation and memory updating as reflected by N170, EPN and P3 amplitudes toward new threat-identities displaying angry expressions. These findings demonstrate that person perception flexibly re-adjusts according to minimal information. Intriguingly, perceptual biases occur even though the anticipated aversive consequence does not occur, with implications for research on stereotyping and anxious psychopathology.
... Taken together, the effects of socially learned aversive anticipations were not reduced by viewing facial images of loved ones, such as one's romantic partner. The knowledge that a particular person or situation is potentially dangerous triggers caution, aversive anticipation, and prepares avoidance or defense behavior 6,8,25,34,35 . Threat-related psychophysiological responding here involves preparatory activation of the somatic and autonomic nervous systems, and is adaptive in that potential harm to the organism can be avoided or reduced. ...
... Indeed, this would be in line with the notion that the human face (compared to pictures of snakes or spiders) 36,37 appears to be a less reliable source of implicit or inherited threat or safety information. For example, social acquisition of threat and safety associations appears to function equally well when linked to facial expressions (e.g., happy or angry faces as threat cues) 34,38 and facial identity information, even for beloved familiar faces 25,35 . Thus, psychophysiological responding reflects a defense mechanism that reacts faithfully to the threatening situation (i.e., colored background), while disregarding and possibly overshadowing prominent foreground information (i.e., pictures of loved ones) that is non-diagnostic with respect to the anticipated danger (i.e., shocks). ...
Article
Full-text available
Looking at pictures of loved ones, such as one’s romantic partner or good friends, has been shown to alleviate the experience of pain and reduce defensive reactions. However, little is known about such modulatory effects on threat and safety learning and the psychophysiological processes involved. Here, we explored the hypothesis that beloved faces serve as implicit safety cues and attenuate the expression of fear responses and/or accelerate extinction learning in a threatening context. Thirty-two participants viewed pictures of their loved ones (romantic partner, parents, and best friend) as well as of unknown individuals within contextual background colors indicating threat-of-shock or safety. Focusing on the extinction of non-reinforced threat associations (no shocks were given), the experiment was repeated on two more test days while the defensive startle-EMG, SCR, and threat ratings were obtained. Results confirmed pronounced defensive responding to instructed threat-of-shock relative to safety context (e.g., threat-enhanced startle reflex and SCR). Moreover, threat-potentiated startle response slowly declined across test days indicating passive extinction learning in the absence of shocks. Importantly, neither a main effect of face category (loved vs. unknown) nor a significant interaction with threat/safety instructions was observed. Thus, a long-term learning history of beneficial relations (e.g., with supportive parents) did not interfere with verbal threat learning and aversive apprehensions. These findings reflect the effects of worries and apprehensions that persist despite the repeated experience of safety and the pictorial presence of loved ones. How to counter such aversive expectations is key to changing mal-adaptive behaviors (e.g., avoidance or stockpiling), biased risk perceptions, and stereotypes.
... While AFC is thought to generate reproducible results for similar faces, human face perception is highly specialized [51] and can be substantially influenced by goals [52], internal states [53], or perceived relevance to the observer [54]. AFC is context independent which is a great advantage in terms of an objective research tool on the one hand. ...
Article
Full-text available
Automatic facial coding (AFC) is a novel research tool to automatically analyze emotional facial expressions. AFC can classify emotional expressions with high accuracy in standardized picture inventories of intensively posed and prototypical expressions. However, classification of facial expressions of untrained study participants is more error prone. This discrepancy requires a direct comparison between these two sources of facial expressions. To this end, 70 untrained participants were asked to express joy, anger, surprise, sadness, disgust, and fear in a typical laboratory setting. Recorded videos were scored with a well-established AFC software (FaceReader, Noldus Information Technology). These were compared with AFC measures of standardized pictures from 70 trained actors (i.e., standardized inventories). We report the probability estimates of specific emotion categories and, in addition, Action Unit (AU) profiles for each emotion. Based on this, we used a novel machine learning approach to determine the relevant AUs for each emotion, separately for both datasets. First, misclassification was more frequent for some emotions of untrained participants. Second, AU intensities were generally lower in pictures of untrained participants compared to standardized pictures for all emotions. Third, although profiles of relevant AU overlapped substantially across the two data sets, there were also substantial differences in their AU profiles. This research provides evidence that the application of AFC is not limited to standardized facial expression inventories but can also be used to code facial expressions of untrained participants in a typical laboratory setting.
... The role of "top down" and "bottom up" mechanisms and their interactions is still debated (Abado, Aue, & Okon-Singer, 2020;Sussman, Jin, & Mohanty, 2016). Various empirical findings suggest the influence of "top down" factors on perception (e.g., Bublatzky, Guerra, & Alpers, 2018;Bublatzky, Guerra, & Alpers, 2020;Kavcioglu, Bublatzky, Pittig, & Alpers, 2021), also specifically for the perception under the condition of binocular rivalry (for a review see Paffen & Alais, 2011). Some research suggest that active voluntary "top down" mechanisms can exert a major influence on perception under binocular rivalry (Leopold & Logothetis, 1999) while other research suggest that the perception under binocular rivalry involves a more automatic, stimulus-driven form of visual competition (Meng & Tong, 2004). ...
Article
What we see is the result of an efficient selection of cues in the visual stream. Not only physical characteristics but also emotional salience influences this process. Previously, we showed in spider phobic patients that fear-related pictures gain preferential access to consciousness in binocular rivalry. We set out to replicate this in an independent unselected sample and examine the relationship of this perceptual bias with a range of symptom clusters. To this end, we recruited 79 participants with variable degrees of fear of spiders. To induce binocular rivalry, a picture of either a spider or a flower was projected to one eye, and a neutral geometric pattern to the other eye. Participants continuously reported what they saw. We correlated indices of perceptual dominance (first percept, dominance duration) with individual fear of spiders and with scores on specific symptom clusters of fear of spiders (i.e., vigilance, fixation, and avoidance coping). Overall, higher fear of spiders correlates with more predominant spider pictures. In addition, this perceptual bias is uniquely associated with avoidance coping. Interestingly, this demonstrates that a perceptual bias, which is not intentionally controlled, is linked with an instrumental coping behavior, which has been implicated in the maintenance of pathological fear.
... Indeed, this would be in line with the notion that the human face (compared to pictures of snakes or spiders; e.g., Lipp & Edwards, 2002;Öhman & Mineka, 2001) appears to be a less reliable source of implicit or inherited threat or safety information. For example, social acquisition of threat and safety associations appears to function equally well when linked to facial expressions (e.g., happy or angry faces as cues to threat; Bublatzky et al., 2018 and facial identity information, even for beloved familiar faces (Bublatzky et al., 2020;Morato et al., 2021). Thus, psychophysiological responding re ects a exible defense activation to the threatening contextual situation (i.e., colored background), disregarding and potentially overshadowing foreground pictures of loved ones or unknown persons that are non-diagnostic with respect to shock threat. ...
Preprint
Full-text available
Looking at pictures of loved ones, such as one's romantic partner or good friends, has been shown to alleviate the experience of pain and reduce defensive reactions. However, little is known about such modulatory effects on threat and safety learning and the psychophysiological processes involved. Here, we explored the hypothesis that beloved faces serve as implicit safety cues and attenuate the expression of fear responses and/or accelerate extinction learning in a threatening context. Thirty-two participants viewed pictures of their loved ones (romantic partner, parents, and best friend) as well as of unknown individuals within contextual background colors indicating threat-of-shock or safety. Focusing on the extinction of non-reinforced threat associations (no shocks were given), the experiment was repeated on two more test days while the defensive startle-EMG, SCR, and threat ratings were obtained. Results confirmed pronounced defensive responding to instructed threat-of-shock relative to safety context (e.g., threat-enhanced startle reflex and SCR). Moreover, threat-potentiated startle response slowly declined across test days indicating passive extinction learning in the absence of shocks. Importantly, neither a main effect of face category (loved vs. unknown) nor a significant interaction with threat/safety instructions was observed. Thus, a long-term learning history of beneficial relations (e.g., with supportive parents) did not interfere with verbal threat learning and aversive apprehensions.
... This threatof-shock paradigm (Grillon et al., 1991) has been shown to reliably trigger selective attention, prime defensive physiological responding, and provoke aversive apprehensions towards instructed threat cues, even without receiving electric shocks (e.g. Bublatzky et al., 2010Bublatzky et al., , 2014Bublatzky et al., , 2018Bublatzky et al., , 2019Costa et al., 2015;Robinson et al., 2013). In a following recognition session, all previously seen (old) faces were presented without background colours intermixed with new unseen faces, and a combined item/source memory task was performed (e.g. ...
Article
To organise future behaviour, it is important to remember both the central and contextual aspects of a situation. We examined the impact of contextual threat or safety, learned through verbal instructions, on face identity recognition. In two studies (N = 140), 72 face–context compounds were presented each once within an encoding session, and an unexpected item/source recognition task was performed afterwards (including 24 new faces). Hierarchical multinomial processing tree modelling served to estimate individual parameters of item (face identity) and source memory (threat or safety context) as well as guessing behaviour. Results show that language was highly effective in establishing threatening and safe context conditions. In Study 1, a fleeting picture stream (1 s per picture) led to poor item and source recognition. Prolonged presentation times (Study 2 with 6 s per picture) improved face memory but no contextual modulation was observed. Thus, incidental face learning was surprisingly poor and rapidly changing contextual settings might have interfered with the accurate encoding of face identity information and item–source binding.
... Lonsdorf et al., 2019;. Conversely, the verbal induction of anticipatory anxiety was more salient than the cue conditioning (see also Bublatzky, Guerra, & Alpers, 2018), potentially explaining why transient SCRs to the central cues were mainly driven by context effects. ...
Thesis
Full-text available
Adapting defensive behavior to the characteristics of a threatening situation is a fundamental function of the brain. Particularly, threat imminence plays a major role for the organization of defensive responses. Acute threat prompts phasic physiological responses, which are usually associated with an intense feeling of fear. In contrast, diffuse and potentially threatening situations elicit a sustained state of anxious apprehension. Detection of the threatening stimulus defines the key event in this framework, initiating the transition from potential to acute threat. Consequently, attention to threat is crucial for supporting defensive behavior. The functions of attention are finely tuned to the characteristics of a threatening situation. Potential threat is associated with hypervigilance, in order to facilitate threat detection. Once a threatening stimulus has been identified, attention is selectively focused on the source of danger. Even though the concepts of selective attention and hypervigilance to threat are well established, evidence for their neural correlates remain scarce. Therefore, a major goal of this thesis is to elucidate the neural correlates of selective attention to acute threat and hypervigilance during potential threat. A second aim of this thesis is to provide a mechanistic account for the interaction of fear and anxiety. While contemporary models view fear and anxiety as mutually exclusive, recent findings for the neural networks of fear and anxiety suggest potential interactions. In four studies, aversive cue conditioning was used to induce acute threat, while context conditioning served as a laboratory model of potential threat. To quantify neural correlates of selective attention and hypervigilance, steady-state visual evoked potentials (ssVEPs) were measured as an index of visuocortical responding. Study 1 compared visuocortical responses to acute and potential threat for high versus low trait-anxious individuals. All individuals demonstrated enhanced electrocortical responses to the central cue in the acute threat condition, suggesting evidence for the neural correlate of selective attention. However, only low anxious individuals revealed facilitated processing of the contexts in the potential threat condition, reflecting a neural correlate of hypervigilance. High anxious individuals did not discriminate among contexts. These findings contribute to the notion of aberrational processing of potential threat for high anxious individuals. Study 2 and 3 realized orthogonal combinations of cue and context conditioning to investigate potential interactions of fear and anxiety. In contrast to Study 1 and 2, Study 3 used verbal instructions to induce potentially threatening contexts. Besides ssVEPs, threat ratings and skin conductance responses (SCRs) were recorded as efferent indices of defensive responding. None of these studies found further evidence for the neural correlates of hypervigilance and selective attention. However, results for ratings and SCRs revealed additive effects of fear and anxiety, suggesting that fear and anxiety are not mutually exclusive, but interact linearly to organize and facilitate defensive behavior. Study 4 tested ssVEPs to more ecologically valid forms of context conditioning, using flickering video stimuli of virtual offices to establish context representations. Contrary to expectations, results revealed decreased visuocortical responses during sustained presentations of anxiety compared to neutral contexts. A disruption of ssVEP signals eventually suggests interferences by continuously changing video streams which are enhanced as a function of motivational relevance. In summary, this thesis provided evidence for the neural correlates of attention only for isolated forms of fear and anxiety, but not for their interaction. In contrast, an additive interaction model of fear and anxiety for measures of defensive responding offers a new perspective on the topography of defensive behavior.
... For instance, enhanced activity of the somatic and autonomic nervous system has been shown for instructed threat relative to safety cues (e.g. enhanced skin conductance responses, heart rate deceleration, and potentiated startle responses; Bradley, Moulder, & Lang, 2005;Bublatzky, Gerdes, White, Riemer, & Alpers, 2014;Bublatzky, Guerra, Pastor, Schupp, & Vila, 2013;Bublatzky, Guerra, & Alpers, 2018;Bublatzky, Riemer, & Guerra, 2019;Costa et al., 2015). Moreover, instructed threat modulates perceptual processing and behavioral responding such as decision-making and memory performance (e.g. ...
Article
Often the source of information is as important as the information itself. The present study examined the impact of contextual threat and safety signals (source information) on memory encoding and recognition of faces (item information). In two experimental sessions, 30 participants viewed neutral face pictures. In the encoding session, 60 faces were presented with contextual background colors (blue or green, 30 pictures each) which were verbally instructed to signal either threat-of-shock or safety. In the recognition session, the 60 old faces intermixed with 30 new faces were shown while a combined old/new recognition and source memory task was performed. During the encoding session, face processing varied as a function of contextual source information. Confirming successful threat-of-shock manipulation, threatening compared to safe face–context compounds revealed differential neural processing (early parieto-occipital and late fronto-central negativity) as well as pronounced threat ratings. During the recognition session, participants had serious problems identifying old from new faces with poor source memory. Intriguingly, however, brain activity differentiated previously seen faces from newly presented pictures (old/new ERP effect). Moreover, old faces presented within a threat context were associated with distributed late negativities compared to old safe faces. Thus, threat effects not only emerged during face encoding (incidental learning) but also during face recognition, although no valid judgements could be made regarding the threatening or safe sources. These findings support the notion that contextual source information critically modulates person perception and recognition as a form of an expectation based remembering in the absence of conscious recognition.
Article
Threat changes cognition and facilitates adaptive coping. However, when threat becomes overwhelming, it may be deleterious to mental health, especially for vulnerable individuals. Flexible decision-making was probed with a reward reversal task to investigate how well healthy participants (N = 34) can adapt to changes in reward contingency when they expect adverse events (i.e., electric shocks). In comparison to a safe control condition, the threat of shock significantly impaired reward reversal learning. Moreover, enhanced self-reported threat ratings and elevated skin conductance levels support the successful induction of stressful and aversive apprehensions. The findings are in line with literature showing the stress-induced inhibition of goal-directed behavior at the advantage of a reflexive (habitual) response style. Notably, reversal learning was rapidly restored with the omission of threat through several cycles of threat and safety contexts within one experimental session. These results extend the literature and illuminate the immediate consequence of a sustained threatening stressor (and its removal) on decision-making. Better knowledge of the immediate effects of anticipatory anxiety on behavior could improve understanding of psychopathology and may be informative for the development of effective therapy for anxiety and emotion dysregulation.
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Ostracism, or social exclusion, is widespread and associated with a range of detrimental psychological and social outcomes. Ostracism is typically explained as instrumental punishment of free-riders or deviants. However, this instrumental account fails to explain many of the features of real-world ostracism, including its prevalence. Here we hypothesized that ostracism can emerge incidentally (non-instrumentally) when people choose partners in social interactions, and that this process is driven by simple learning mechanisms. We tested this hypothesis in four experiments (n = 456) with economic games in dynamic social networks. Contrary to the instrumental account of ostracism, we find that the targets of ostracism are not primarily free-riders. Instead, incidental initial variability in choosing partners for social interactions predicts later ostracism better than the instrumental account. Using computational modelling, we show that simple reinforcement learning mechanisms explain the incidental emergence of ostracism, and that they do so better than a formalization of the instrumental account. Finally, we leveraged these reinforcement learning mechanisms to experimentally reduce incidental ostracism. Our results demonstrate that ostracism is more incidental than previously assumed and can arise from basic forms of learning. They also show that the same mechanisms that result in incidental ostracism can help to reduce its emergence.
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