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The Silences of the Archives, the Reknown of the Story. The Martin Guerre affair has been told many times since Jean de Coras and Guillaume Lesueur published their stories in 1561. It is in many ways a perfect intrigue with uncanny resemblance, persuasive deception and a surprizing end when the two Martin stood face to face, memory to memory, before captivated judges and a guilty feeling Bertrande de Rols. The historian wanted to go beyond the known story in order to discover the world of the heroes. This research led to disappointments and surprizes as documents were discovered concerning the environment of Artigat’s inhabitants and bearing directly on the main characters thanks to notarial contracts. Along the way, study of the works of Coras and Lesueur took a new direction. Coming back to the affair a quarter century later did not result in finding new documents (some are perhaps still buried in Spanish archives), but by going back over her tracks, the historian could only be struck by the silences of the archives that refuse to reveal their secrets and, at the same time, by the possible openings they suggest, by the intuition that almost invisible threads link here and there characters and events.
Heartfelt imitation: high interoceptive 1
awareness is linked to greater automatic 2
imitation. 3
Vivien Ainley1, Marcel Brass2, & Manos Tsakiris1 5
1Lab of Action & Body, Department of Psychology, Royal Holloway, University of 9
London, UK 10
2Department of Experimental Psychology, Ghent University 11
Corresponding author: Vivien Ainley, Department of Psychology, Royal Holloway, 13
University of London, Egham, Surrey, UK. Tel. +44(0)1784276551, Fax. 14
+44(0)1784434347, E-mail: 15
Funding: This study was funded by the European Platform for Life Sciences, Mind 18
Sciences and Humanities, Volkswagen Foundation (II/85 064) and European 19
Research Council Starting Investigator Grant (ERC-2010-StG-262853) to MT20
Abstract 21
‘Interoceptive awareness’, defined as the individual’s awareness of internal body 22
signals, modulates self/other distinction under conditions of multisensory integration. 23
We examined here, for the first time, the potential impact of interoceptive awareness 24
on self/other distinction in the motor domain. In automatic imitation, inhibition of 25
imitation is an index of an individual’s success in distinguishing internally generated 26
motor representations from those triggered by observing another person’s action. This 27
is measured by the ‘congruency effect’, which is the difference between mean 28
reaction times when the observed action is ‘incongruent’ with the required action and 29
when it is ‘congruent’. The present study compared the congruency effect in a typical 30
finger lifting paradigm, with interoceptive awareness measured by heartbeat 31
perception. Contrary to expectation, interoceptive awareness was positively correlated 32
with the congruency effect and this effect depended on mean reaction times in the 33
incongruent condition, indicating that good heartbeat perceivers had more difficulty 34
inhibiting the tendency to imitate. Potentially, high interoceptive awareness involves 35
stronger interoceptive representations of the consequences of an action, implying 36
higher empathy, greater motor reactivity in response to observed action and hence a 37
greater tendency to imitate. Our results may also tentatively be explained within a 38
predictive coding account of interoception. 39
Highlights 43
Interoceptive awareness modulates self/other distinctions in body-awareness 44
tasks. 45
Automatic imitation also indexes the ability to distinguish ‘self’ from ‘other’. 46
In a finger-lifting task, good heartbeat perceivers had larger ‘congruency 47
effects’. 48
Interoceptive awareness correlated with difficulty in inhibiting imitation. 49
1. Introduction 51
The ability to distinguish between self and other is crucial to all aspects of self-52
processing and has relevance for action-awareness (Farrer et al., 2003), body-53
awareness (Tsakiris, 2013), empathy (Singer et al., 2004) and social cognition 54
(Lamm, Batson, & Decety, 2007). In the motor domain, self/other distinction has been 55
extensively studied using ‘automatic imitation’ paradigms (Brass, Bekkering, & Prinz, 56
2001; Catmur, Walsh, & Heyes, 2007), where the ability to resist imitating an action 57
performed by another person is taken to indicate a stronger sense of self (Spengler, 58
Brass, Kühn, & Schütz-Bosbach, 2010). Recent theories propose, however, that the 59
self is grounded in ‘interoception’, which refers to the signals arising from within the 60
body (Craig, 2010; Damasio, 2010; Seth, 2013). Awareness of such internal signals 61
has been shown to influence the ability to distinguish between self and other in 62
multisensory contexts (Suzuki, Garfinkel, Critchley, & Seth, 2013; Tsakiris, Tajadura-63
Jiménez, & Costantini, 2011). Given the inter-connectedness of perception and action 64
(Friston, 2010; Hommel, 2009) the purpose of this study was to investigate whether 65
awareness of interoceptive cues similarly impacts on self/other distinction in the 66
domain of action. 67
Humans have a tendency to involuntarily imitate actions that they observe. Thus, 69
when an individual is required to perform a given action, observing another person 70
perform an identical action typically facilitates performance, whereas observing a 71
different action generally interferes with it, even when the observed action is entirely 72
task-irrelevant (see Heyes, 2010, for a review). Although the term ‘automatic 73
imitation’ is commonly used, the phenomenon rarely involves true imitation, in that 74
people actually seldom perform the wrong action. They must, however, resist a 75
tendency to copy the action they observe. The ability to inhibit imitation is measured 76
by ‘the congruency effect’, which is the difference between the slower mean reaction 77
time (RT) typically found when the required and observed actions are ‘incongruent’ 78
(i.e. different) and the faster mean RT when the desired and observed actions are 79
‘congruent’ (Brass, Bekkering, Wohlschläger, & Prinz, 2000). 80
According to the Theory of Event Coding, automatic imitation occurs because actions 82
are coded in terms of their goals and thus their sensory consequences. The distinction 83
between perception and action is thus a false dichotomy (Hommel, Müsseler, 84
Aschersleben, & Prinz, 2001) and seeing an action necessarily primes the motor 85
representation of that action. The Associative Sequence Learning (ASL) theory 86
(Catmur, Walsh, & Heyes, 2009), suggests that visual and motor components of 87
actions are linked by long-term stimulus response (SR) bonds, such that the activation 88
of a visual mental representation necessarily predicts a motor representation (Heyes, 89
2010). More recently, the theory of predictive coding has linked perception and action 90
within a unified framework that may, in future, elucidate the neural mechanisms 91
behind automatic imitation (Adams, Shipp, & Friston, 2012; Friston, 2010). 92
Not only does automatic imitation rarely involve imitation but neither is it truly 94
‘automatic’, because it is not immune to interference by other processes. According to 95
the ASL model (Catmur et al., 2009) these processes can be divided into ‘input 96
modulation’, which alters the extent to which the relevant long-term SR bond is 97
activated, and ‘output modulation’, where social factors potentially inhibit the 98
involuntary imitation (Heyes, 2010). Input modulation is demonstrated by selective 99
attention to one’s own actions, which reduces imitation (Bortoletto, Mattingley, & 100
Cunnington, 2013; Chong, Cunnington, Williams, & Mattingley, 2009). Automatic 101
imitation also can be reduced by modest amounts of training (Cook, Press, Dickinson, 102
& Heyes, 2010; Gillmeister, Catmur, Brass, & Heyes, 2008; Heyes, Bird, Johnson, & 103
Haggard, 2005; Heyes & Bird, 2007), which reverses the muscle specificity of the 104
motor-evoked potentials (MEPs) produced by TMS (Catmur et al., 2007). 105
Output modulation depends on the top-down influence of participants’ traits and 107
social attitudes. Eye contact, or priming with pro-social cues, enhances the 108
congruency effect (Leighton, Bird, Orsini, & Heyes, 2010; Wang & Hamilton, 2012; 109
Wang, Newport, & Hamilton, 2011). Similarly, a desire to affiliate to the person 110
observed increases automatic imitation in both experimental settings and social 111
interaction (Lakin & Chartrand, 2003; Wang & Hamilton, 2012). People scoring high 112
in ‘self-monitoring’ (Snyder, 1974), or who have an interdependent self-construal, 113
have a greater tendency to mimic others, possibly as an unconscious affiliation 114
strategy (Cheng & Chartrand, 2003; Obhi, Hogeveen, & Pascual-Leone, 2011). 115
Interestingly, priming participants with examples of interdependent self-construal 116
increases the amplitude of MEPs elicited by TMS (Obhi et al., 2011), indicating that 117
these top-down influences increase cortical excitability in the motor areas that 118
produce imitation. 119
Automatic imitation is one of a number of phenomena which involve ‘self/other 121
overlap’, defined as “any phenomenon whereby an observer engages a state similar 122
to that of the target, via activation of the observer’s personal representations for 123
experiencing the observed state, whether through direct perception or simulation” 124
(Preston & Hofelich, 2012). These shared representations occur at a very early, 125
preconscious, processing stage. The ability to inhibit imitation requires that the 126
individual distinguishes between internally generated motor representations and those 127
that are triggered by observing other people’s actions (Brass, Ruby, & Spengler, 128
2009). Successfully inhibiting the tendency to imitate activates cortical areas thought 129
to be involved in discriminating between self and other (Brass, Derrfuss, & von 130
Cramon, 2005; Brass et al., 2009; Brass & Heyes, 2005). The most active of these 131
regions - the temporal parietal junction and anterior fronto-median cortex (BA10) - 132
are related to perspective taking, feelings of agency and theory of mind (Wang, 133
Ramsey, & Hamilton, 2011). Greater activation in BA10 correlates with smaller 134
congruency effects and thus with better self/other distinction (Spengler et al., 2009). 135
Furthermore, experimentally increasing self-focus reduces the congruency effect, by 136
reducing RTs on incongruent trials (Spengler, Brass, Kühn, & Schütz-Bosbach, 137
2010). Similarly, observing an action increases the amplitude of MEPs if that action is 138
attributed to another individual but reduces cortico-spinal excitability when the action 139
is illusorily attributed to the self (Schutz-Bosbach, Mancini, Aglioti, & Haggard, 140
2006). 141
Automatic imitation can therefore be characterised as a tool to measure how 143
effectively the self can be distinguished from others (Spengler, von Cramon, & Brass, 144
2009). The purpose of the current experiment was to investigate how the congruency 145
effect is linked to ‘interoceptive awareness’ - a fundamental dimension of self-146
awareness that has been the focus of recent research in body ownership (Tsakiris et 147
al., 2011), self-recognition (Tajadura-Jiménez & Tsakiris, 2013) and empathy 148
(Fukushima, Terasawa, & Umeda, 2011). 149
Recent neuroscientific models of the self emphasize the role of ‘interoception’ (Craig, 151
2010; Critchley & Harrison, 2013; Hayes & Northoff, 2012; Panksepp & Northoff, 152
2009) defined as “the afferent information arising from within the body, affecting the 153
cognition, emotion or behaviour of an organism, with or without awareness” 154
(Cameron, 2001). Insular cortex, which is activated by all feelings arising within the 155
body (Craig, 2010; Critchley & Harrison, 2013; Singer, Critchley, & Preuschoff, 156
2009; Wiebking et al., 2013; Zaki, Davis, & Ochsner, 2012), may underpin this 157
fundamental representation of self (Craig, 2009; Seth, 2013; but see also Philippi et 158
al., 2012). Recent predictive coding accounts of cortical function (Clark, 2013; 159
Friston, 2010) similarly propose interoceptive information as an essential component 160
of the self (Apps & Tsakiris, 2013; Seth, Suzuki, & Critchley, 2011). ‘Interoceptive 161
awareness’, which is the extent to which internal signals reach consciousness, has 162
been extensively studied in relation to emotion, stemming originally from William 163
James’ theory that emotion comprises unconscious bodily responses (Damasio & 164
Carvalho, 2013; James, 1890). 165
Recent studies have begun to investigate the contribution of interoceptive awareness 167
to self-processing. In the rubber hand illusion, people with low interoceptive 168
awareness are more likely to claim ownership over a prosthetic hand, (Tsakiris et al., 169
2011) and similarly experience a stronger illusory identification with a stranger’s face 170
when they observe that face being stroked synchronously with felt touch on their own 171
face (Tajadura-Jiménez & Tsakiris, 2013). Conversely, enhanced self-focus, through 172
mirror self-observation, a self-photograph or self-relevant words, can improve 173
interoceptive awareness in people for whom this is initially low (Ainley, Maister, 174
Brokfeld, Farmer, & Tsakiris, 2013; Ainley, Tajadura-Jiménez, Fotopoulou, & 175
Tsakiris, 2012; Maister, Tsiakkas, & Tsakiris, 2013). Individuals who see a virtual 176
image of their own hand (Suzuki, Garfinkel, Critchley, & Seth, 2013) or of their 177
whole body (Aspell et al., 2013) have a greater sense of self-identification with, and 178
self-location towards, the image under conditions of cardio-visual synchrony. 179
Despite these investigations into the contribution of interoceptive awareness to 181
self/other distinction in multisensory contexts, little is known about the potential role 182
of interoception in the action system, for example in automatic imitation. This lack of 183
empirical research is striking, given that human actions are thought to be driven by 184
the goal of homeostatic control, which is signaled interoceptively (Craig, 2010; 185
Damasio, 2010; Seth, 2013). Theoretical accounts of the neural basis of perception 186
and action stress their inter-connectedness (Friston, 2010; Schütz-Bosbach & Prinz, 187
2007). While it has been previously assumed that the sensory consequences of an 188
action are primarily exteroceptive, empathy for pain (Avenanti, Bueti, Galati, & 189
Aglioti, 2005; Singer et al., 2004) and overlapping cortical activation during the 190
experience, observation or imagination of disgust (Wicker et al., 2003) can only be 191
explained if actions involve a representation of their interoceptive sensory 192
consequences (Heyes & Bird, 2007). 193
Given that the ability to inhibit automatic imitation seems to index better self/other 195
distinction, at the level of visual and motor representation, and also that people with 196
high interoceptive awareness appear more reliably able to distinguish their own 197
bodies from those of others, at a multisensory level, we hypothesised that in an 198
automatic imitation paradigm individuals with high interoceptive awareness would 199
successfully inhibit the tendency to imitate, whereas those with low interoceptive 200
awareness would exhibit less self/other distinction and would therefore have a greater 201
tendency to automatic imitation. 202
‘Interoceptive awareness’ is generally assessed using a heartbeat perception task 204
(Schandry, 1981; Whitehead & Drescher, 1980). Such measures correlate with 205
awareness of gastric cues (Herbert, Muth, Pollatos, & Herbert, 2012; Whitehead & 206
Drescher, 1980). We used the Mental Tracking task (Schandry, 1981) which is well-207
validated (Knoll & Hodapp, 1992), with good test retest reliability (Mussgay, 208
Klinkenberg, & Rüddel, 1999; Werner, Kerschreiter, Kindermann, & Duschek, 2013) 209
and which discriminates well between individuals. The measure we have called 210
‘interoceptive awareness’ in this study assesses the accuracy of cardiac awareness, by 211
comparing the subjectively reported number of heartbeats experienced with the 212
number (objectively) recorded (Cuenen, Van Diest, & Vlaeyen, 2012; Garfinkel & 213
Critchley, 2013). Gender, body mass index (BMI), and resting heart rate were also 214
recorded, as possible confounds of the heartbeat perception task (Cameron, 2002). 215
Automatic imitation was assessed using an established inhibition imitation paradigm 216
developed by Brass and colleagues (Brass et al., 2005; Spengler et al., 2009). It was 217
anticipated that people who performed accurately in heartbeat perception would also 218
be more accurate during the automatic attention task (show a smaller congruency 219
effect). However, both these variables might be affected by participants’ general 220
willingness and ability to attend to the tests. Attention is a possible source of input 221
modulation in automatic imitation (Davis, 1983; Kaplan & Iacoboni, 2006; Preston & 222
Hofelich, 2012). It has also been reported (Matthias, Schandry, Duschek, & Pollatos, 223
2009) that interoceptive awareness is linked to scores on the d2 test (Brickenkamp & 224
Zilmer, 1998), which measures individual differences in motivation and attention. We 225
accordingly administered the d2 test as a check for this potential confound. 226
2. Method 228
2.1 Ethics statement 229
The study was approved by the Department of Psychology Ethics Committee, Royal 230
Holloway University of London. All participants gave written informed consent and 231
were free to withdraw from the experiment at will. 232
2.2 Participants 234
Participants were 45 students at Royal Holloway University of London who 235
participated for course credit. All declared themselves right handed and had normal or 236
corrected to normal vision. The data for 2 participants was excluded for excessive 237
numbers of errors (more than 10%, i.e. 3SD above the mean) in the action imitation 238
task, indicating a failure to concentrate and follow the instructions. Of the remaining 239
43 participants, mean age = 19.6 (SD = 4.9), 9 were male. 240
2.3 Stimuli 242
The stimuli consisted of sequences of 5 frames (Brass et al., 2005; Spengler et al., 243
2009). Each video stared with a frame showing the hand, which mirrored the right 244
hand of the subject, in the starting position, for 2s. The next two frames, each lasting 245
34ms, presented a number (either 1 or 2) and simultaneously showed the finger 246
movement (if any). The fourth frame showed the finger in the end position for 1.3s, 247
with the number (1 or 2) superimposed. Between trials, the screen turned black for 248
2.7s. Each video trial was thus 6s duration. The video hand was presented on a blue 249
rectangular background, measuring 22 x 12cm. 250
Figure 1. Example of the video stimuli for the index finger, from Brass et al., (2005) 252
There were six possible video sequences, consisting of each of the two fingers (index 255
or middle) in each of three conditions (baseline, congruent or incongruent). 256
Participants were required to lift either the index (1) or middle (2) finger in response 257
to a number appearing on the screen. The three possible conditions (for the index 258
finger) are shown in Figure 1. Thus in the baseline condition, simultaneous with the 259
appearance of the number, the video hand remained static. In the congruent condition 260
the video hand lifted the finger that corresponded to the number shown (i.e. the index 261
finger was lifted when the number 1 appeared). In the incongruent condition the video 262
hand lifted the ‘wrong’ finger (i.e. the middle finger was lifted when the number 1 263
appeared). 264
2.4 Procedure 266
2.4.1 Interoceptive awareness 267
After giving informed consent, participants’ gender, age, height and weight were 268
recorded. Heartbeat signals were acquired with a piezo-electric pulse transducer, fitted 269
to the participant’s left index finger and connected to a physiological data unit (26T 270
PowerLab, AD Instruments) sampling at 1 kHz which recorded the derived electrical 271
signal onto a second PC running LabChart6 software (AD Instruments). Instructions 272
for the Mental Tracking Method (Schandry, 1981) were presented over noise-273
attenuating headphones. The onset and offset of each heartbeat counting trial were 274
cued by the words “go” and “stop”, presented audiovisually. We used a standard 275
instruction (Ehlers, Breuer, Dohn, & Fiegenbaum, 1995) whereby participants were 276
asked to concentrate hard and try to silently count their own heartbeats, simply by 277
“listening” to their bodies, without taking their pulse. The three trials (25s, 35s & 45s) 278
were presented in random order. A criticism of the Mental Tracking Method is that 279
participants may estimate the elapsed interval and then use knowledge of their own 280
heart rate to guess the number of heartbeats. We therefore asked individuals to 281
estimate the length of three, randomly presented, intervals (19s, 37s, 49s) and to 282
provide an estimate of their resting heart rate (Dunn et al., 2010). 283
2.4.2 Action imitation 285
The stimuli were viewed on a standard PC, using Presentation software 286
(Neurobehavioral Systems, Albany, CA). Participants were seated about 60 cm in 287
front of the screen and were instructed to execute their movements as quickly and 288
accurately as possible. Participants placed the index and middle fingers of their right 289
hand on a serial response box which was linked to another PC which recorded the 290
times of all finger movements, using Spike2 software (Cambridge Electronic Design, 291
Cambridge UK). This recorded the onset of the visible stimulus on screen (i.e. the 292
number 1 or 2, which coincided with the onset of movement of the video hand) and 293
also recorded whenever the participant lifted an index or middle finger. Following 6 294
tests trials, 150 trials experimental trials were presented in three blocks of 5mins, with 295
obligatory rests of at least 2mins between blocks. The order of the presentation of the 296
trials was fully randomised and comprised 25 trials in each of the 6 conditions. 297
2.4.3 The d2 test of attention 299
Finally, the d2 test was administered (Brickenkamp & Zilmer, 1998). This is a widely 300
used measure of selective visual attention. The test items consist of the letters d and p 301
with up to four dashes, arranged either individually or in pairs, above and/or below 302
each letter. The subject is given 20s to scan across each of the 14 closely printed test 303
lines, during which they must identify and cross out every letter d which has exactly 304
two dashes, while ignoring all other distractor letters. The d2 test produces several 305
norm-referenced scores, of which the most commonly reported are the total number of 306
items processed (TN) regardless of whether these are correct or incorrect (this is a 307
measure of processing speed), the percentage of errors made (E%) and the total 308
number of items processed correctly (TN-E). This final score is designed to provide a 309
measure of the capacity to selectively orient to relevant aspects of the task, while 310
screening out irrelevant ones. 311
2.5 Data reduction 313
2.5.1 Interoceptive Awareness 314
LabChart6 was employed to identify and count the number of R-wave peaks on the 315
heart trace recorded for each participant in each trial, as well as to calculate the 316
average heart rates for each trial (Jennings et al., 1981). Every heart trace was visually 317
inspected for artefacts and the number of R-wave peaks was recounted manually, if 318
necessary. No participant was excluded due to artefacts. Interoceptive awareness was 319
calculated as (1/3Σ (1-(|recorded heartbeats – counted heartbeats|/recorded 320
heartbeats)) (Schandry, 1981). Higher scores indicate higher interoceptive awareness. 321
As a control on guessing, the participant’s ability to estimate the length of an elapsed 322
interval was also calculated as (1/3Σ (1-(|estimated elapsed time – actual elapsed 323
time|/actual elapsed time)) which we called the “time modulus” measure (Dunn et al., 324
2010). 325
2.5.2 Action imitation 327
Data was extracted using Matlab ( and analysed with Microsoft 328
Excel. The mean reaction time (RT) was calculated for each of the 6 conditions 329
(congruent, incongruent and baseline, for each of the two fingers). The ‘congruency 330
effect’ was found by subtracting the mean RT for congruent trials from the mean RT 331
for incongruent trials. 332
3. Results 334
3.1 Error analysis 335
RT errors were removed before analysis. There were 2 possible sources of RT errors. 336
Firstly, participants occasionally lifted the wrong finger. Secondly, in common with 337
most RT analyses, some response times were omitted as outliers (Miller & Diego, 338
1991). Thus RTs less than 80ms or greater than 800ms were excluded from the RT 339
analysis (Brass, Bekkering, & Prinz, 2001). The rate for all errors was 2.3% of trials. 340
Two participants were excluded for total errors > 10% i.e. 3SD above the mean. The 341
distribution of errors was thereafter approximately Normal, skewness = .64, kurtosis = 342
-.16. 343
Paired sample t tests (with Bonferroni correction for multiple comparisons and a 345
significance level of 0.017) showed that there were significantly more errors in the 346
incongruent condition than in the baseline, t(42) = 5.07, p < .001, but no significant 347
difference between the numbers of errors in the congruent condition and baseline, 348
t(42) = 0.82, p = .42, replicating the finding of Brass et al. (2005). 349
3.2 Reaction Time (RT) Analysis 351
Repeated measures ANOVA was performed, with both the finger (index or middle) 352
and the condition (congruent, incongruent and baseline) as within-subjects variables. 353
Mauchley’s test of Sphericity was significant; therefore Greenhouse Geisser 354
corrections were applied. There was a main effect of condition (RTs in the 355
incongruent conditions were slower), F(2, 84) = 186.4, p < .001. This indicates 356
significant automatic imitation i.e. slower mean RTs in the incongruent than 357
congruent condition, for both fingers (Brass et al., 2000; Brass, Derrfuss, & von 358
Cramon, 2005). There was a main effect of finger, F(1, 42) = 13.2, p = .001 (reaction 359
times were generally faster for the middle finger), as shown in Figure 2. The 360
interaction of finger and condition was also significant, F(2, 84) = 8.9, p < .001. 361
Paired samples t tests (with Bonferroni correction and a significance level of 0.008) 362
showed that, compared with RTs in the baseline, RTs in the incongruent condition 363
were significantly longer when participants were required to lift their index finger 364
rather than their middle finger, t(42) = 3.32, p = .002. However, there was no 365
significant difference between the two fingers for RTs in the congruent condition, 366
compared with the baseline, t(42) = .57, p = .57. Despite the significantly shorter RTs 367
for the middle finger, particularly in the incongruent condition, the relationships 368
between interoceptive awareness and the various reaction time measures in our study 369
were very similar for the two fingers. For the remaining analysis we therefore used 370
the mean of the data for the index and middle fingers, to give a single measure of 371
average RT in each condition. 372
Figure 2. Mean reaction times by condition and by finger (Errors bars = SEM) 374
To investigate the relationship between interoceptive awareness and the congruency 377
effect, we calculated the latter, in the standard way (as the mean RT in the 378
incongruent condition minus the mean RT in the congruent condition), for the average 379
of the two fingers, for each participant. Correlations between interoceptive awareness 380
and differences in RTs between conditions are shown in Table 1. Interoceptive 381
awareness was positively correlated with the congruency effect (Figure 3) and this 382
was wholly accounted for by RTs in the incongruent condition. Interoceptive 383
awareness was significantly correlated with the difference between mean RTs in the 384
incongruent condition and the baseline but not with the difference between mean RTs 385
in the congruent and baseline conditions. 386
Figure 3. Scatter diagram of the average congruency effect against interoceptive 388
awareness 389
Table 1. Correlations between interoceptive awareness (IA) and RT measures 393
IA & ‘the congruency effect’ (mean RT in incongruent condition
minus the congruent condition)
r = .41
p = .006**
IA & mean RT in the incongruent condition minus the baseline
r = .45
p = .002**
IA & mean RT in the congruent condition minus the baseline
r = -.04
p = .73
** significant at the 1% level 395
The wide range of mean RTs amongst our participants (318ms - 513ms, median 397
398ms) might have affected our results. We therefore calculated the percentage 398
difference in RTs between the incongruent and congruent conditions using the 399
formula [{(mean RT incongruent - mean RT congruent)/mean RT baseline} x 100]. 400
This statistic was also significantly positively correlated with interoceptive awareness, 401
r = .40, p = .008. 402
In this experiment we recorded a number of confounding variables known to impact 404
on interoceptive awareness, namely gender, Body Mass Index (BMI), resting heart 405
rate, and two measures designed to assess possible guessing on the Mental Tracking 406
task (i.e. the ‘time modulus’ measure of the participant’s ability to estimate elapsed 407
time, and the participant’s belief about his/her heart rate). An independent samples t 408
test (with equal variances not assumed) showed no effect of gender on interoceptive 409
awareness, t(41) = 1.32, p = .24. Likewise the correlation of interoceptive awareness 410
and BMI was not significant, r = -.20, p = .21. Although people with slower hearts are 411
often better heartbeat perceivers (Ainley et al., 2012; Cameron, 2001; Knapp-Kline & 412
Kline, 2005), in this sample the correlation of interoceptive awareness and average 413
heart rate did not reach significance r = -.22, p = .16. 414
The ‘time modulus’ measure (of participants’ ability to estimate the length of an 416
elapsed interval) was correlated with interoceptive awareness, r = .35, p = .02 but the 417
correlation of interoceptive awareness and participants’ estimates of their own heart 418
rates was not significant, r = -.08, p = .62. 419
Table 2. Hierarchical multiple regression with the average congruency effect as the 421
dependent variable 422
Step 1
Step 3
Step 4
Step 5
Step 6
awareness (IA)
(p = .36)
(p =.16)
(p = .14)
0.40 **
(p = .009)
(p = .006)
Average heart
rate (HR)
(p = .82)
‘Time modulus’
(p = .64)
(p = .29)
* significant at the 5% level 423
** significant at the 1% level 424
Given previous correlations in the literature between interoceptive awareness and both 426
participants’ average heart rates and the ‘time modulus’ measure (Cameron, 2002; 427
Dunn et al., 2010), we performed a hierarchical multiple regression analysis with the 428
average congruency effect as the dependent variable and independent variables 429
comprising interoceptive awareness, average heart rate, ‘time modulus’, and their 430
interactions. Only interoceptive awareness had any significant effect on the 431
congruency effect (see Table 2). 432
Results of the d2 test of attention were analysed in terms of the total number of items 434
processed (TN), total number correct (TN-E) and percentage of errors (E%). 435
Compared with published norms, d2 scores for our participants (mean TN = 516, 436
mean (TN-E) = 493) were at the 70th percentile for students. Previous research 437
(Matthias et al., 2009) found significant correlation between interoceptive awareness 438
and TN but in this experiment none of the d2 measures were correlated with 439
interoceptive awareness, for TN r = .03, p = .87, for (TN-E) r = .04, p = .82 and for 440
(E%) r = -.02, p = .92. To replicate the analysis of Matthias et al. (2009), we split the 441
data using their cut off at interoceptive awareness = .85 but found no significant 442
difference in any d2 measures between ‘good’ (interoceptive awareness > .85, n = 5) 443
and ‘poor’ (interoceptive awareness < .85, n = 38) heartbeat perceivers (e.g. for TN, 444
F(1, 41) = .46, p = .50). There were likewise no significant correlations between any 445
of the d2 measures and the average congruency effect, for TN r = .18, p = .24, for 446
(TN-E) r = .11, p = .47, and for (E%) r = .15, p = .32. 447
4. Discussion 449
We investigated the relationship between interoceptive awareness and automatic 450
imitation, measuring interoceptive awareness (IA) with a well-validated heartbeat 451
perception task (Schandry, 1981) and automatic imitation by a widely used finger-452
lifting paradigm (Brass et al., 2005). The expected ‘congruency effect’ was obtained, 453
i.e. mean reaction times (RTs) were slower when the observed and required actions 454
were incongruent and were faster when they were congruent (compared with the 455
baseline of no observed movement). Interoceptive awareness was significantly 456
positively correlated with the congruency effect. This was fully accounted for by the 457
difference between RTs in the incongruent condition and the baseline. There were no 458
Interaction of
IA & ‘time
(p = .86)
Interaction of
(p = .31)
(p = .26)
(p = .69)
Interaction of
‘time modulus’
& HR
(p = .62)
(p = .23)
(p = .40)
(p = .45)
Adjusted R2
(p = .18)
(p = .06)
(p = .05)
(p = .02)
(p = .006)
significant effects of interoceptive awareness on RTs difference between the 459
congruent and the baseline. Thus the relationship we observed depended on RTs the 460
incongruent condition and thus on interference between the observed and required 461
action (Blakemore & Frith, 2005), indicating that people with high interoceptive 462
awareness had greater difficulty inhibiting the tendency to automatically imitate. Had 463
there been a motor facilitation effect, it would have taken the form of shorter RTs on 464
congruent trials. RTs in the incongruent condition were significantly slower for the 465
index finger than for the middle finger, probably because lifting an index finger is a 466
more familiar experience than the isolated lifting of a middle finger, with a 467
consequently stronger, learned associative bond. 468
The result we obtained was contrary to our original hypothesis. Experiments in 470
multisensory integration have suggested that people with high interoceptive 471
awareness are better at making self/other body ownership distinctions (Tajadura-472
Jiménez & Tsakiris, 2013; Tsakiris et al., 2011). We hypothesized that this effect 473
might translate into the motor domain. The ability to inhibit imitation is assumed to 474
index self/other distinction (Spengler et al., 2009) and we therefore predicted that 475
people with high interoceptive awareness would more successfully inhibit the 476
tendency to imitate. Our results show that, on the contrary, they were more inclined to 477
imitate, implying greater self/other overlap. 478
Despite the findings from body-ownership paradigms, which suggest that high 480
interoceptive awareness is linked to better ability to make self/other distinctions, this 481
is likely to be context dependent. Thus while low interoceptive awareness might 482
predict greater ability to distinguish between self and other in cases of multisensory 483
body-related integration (Tajadura-Jiménez & Tsakiris, 2013; Tsakiris et al., 2011), in 484
other contexts high interoceptive awareness seems to suggest greater self/other 485
overlap. A fundamental difference between self/other distinction in the automatic 486
imitation task and self/other distinction in the rubber hand illusion is that confusion in 487
the automatic imitation task is at a representational level and at a point in time where 488
participants have no sensory information about their own movements. The link 489
between interoceptive awareness and automatic imitation may therefore be indirect 490
and depend on the sensitivity of people with high interoceptive awareness to social 491
influences. Thus the concentration of our effect in incongruent cues indicates that it 492
depended on the action observation aspect of the task and therefore on output 493
modulation, rather than the preparation of the individual’s own action (input 494
modulation). The lack of correlation between the congruency effect and the d2 test 495
also supports this conclusion. The d2 test scores are measures of “the capacity to 496
selectively orient to relevant aspects of the task while screening out irrelevant ones” 497
(Zimmerman & Frimm, 2002). The d2 was included to counter the criticism that if we 498
had found the hypothesised correlation between high accuracy in both the heartbeat 499
detection and the automatic imitation tasks, this might have reflected the participants’ 500
level of motivation and attention. We did not replicate previous reports of a 501
correlation between high interoceptive awareness and selective and divided attention 502
(Matthias et al., 2009), indicating that general differences in individuals’ motivation 503
and attention to the tasks were unlikely to have confounded our results. 504
In terms of the Associative Sequence Learning model of automatic imitation (Catmur 506
et al., 2009) output modulation is occasioned by social factors which influence 507
individuals to suppress or enhance the tendency to imitate. High interoceptive 508
awareness has been linked to anxiety (Domschke et al., 2010) and particularly to 509
social anxiety (Terasawa, Shibata, Moriguchi, & Umeda, 2013). We did not assess 510
trait anxiety in this study but potentially, if our high interoceptive awareness 511
participants were more socially anxious, they might have had a greater desire to 512
affiliate, which could have enhanced their tendency to imitate. 513
A potential source of output modulation is affective empathy, which is assumed to 515
involve shared representations between one’s own emotional state and that of another 516
individual (Decety & Jackson, 2004; Iacoboni, 2009; Preston & Hofelich, 2012; Zaki, 517
Weber, Bolger, & Ochsner, 2009). People with high interoceptive awareness are 518
thought to exhibit greater empathy (Ernst et al., 2013; Terasawa, Shibata, Moriguchi, 519
& Umeda, 2013), perhaps because they have a stronger interoceptive representation of 520
the consequences of an observed action, for example, they are more sensitive to 521
masked fear conditioning (Katkin, Wiens, & Ohman, 2001). Scores on the empathetic 522
concern scale of the Interpersonal Reactivity Index (Davis, 1983) correlate with the 523
amplitude of heartbeat evoked potentials (Fukushima, Terasawa, & Umeda, 2011), 524
which are larger in people with high interoceptive awareness (Pollatos & Schandry, 525
2004). Empathy has, in turn, been linked to action observation. Kaplan and Iacoboni 526
(2006) found that when participants observed another individual reaching for a cup, 527
inferior frontal mirror activity was greater in those people who had higher scores on 528
the Empathetic Concern subscale. Such motor activity in response to action 529
observation is also linked to a greater tendency to imitate (Catmur et al., 2007; Obhi 530
et al., 2011; Schutz-Bosbach et al., 2006). Empathy is inversely correlated with 531
narcissism and it has recently been shown that individuals who are high in trait 532
narcissism - thus displaying a lack of empathy and concern for others - have a greater 533
ability to inhibit automatic imitation (Obhi, Hogeveen, Giacomin, & Jordan, 2013). 534
Thus high interoceptive awareness may involve stronger interoceptive representation 535
of the consequences of an action, implying higher empathy, greater mirror neuron 536
activity in response to observed action and hence a greater tendency to imitate. 537
Our results may alternatively depend on some hitherto unexplored aspect of 539
interoceptive awareness and its relationship to the action system. Given that accounts 540
of cortical function, including both the Theory of Event Coding (Hommel, 2009) and 541
predictive coding (Clark, 2013; Friston, 2010) stress that perception and action are 542
reciprocally connected, further research is needed to confirm whether interoceptive 543
awareness impacts not only on action in interoceptive systems but on motor activity 544
as well. The basis of inter-individual differences in interoceptive awareness is not 545
well understood (Verdejo-Garcia, Clark, & Dunn, 2012). Such differences have 546
generally been assumed to depend simply on the strength of interoceptive signals 547
arising within the body, which are conveyed principally by the vagus nerve (Craig, 548
2003; Cameron, 2002; Critchley et al., 2007). However, interoceptive awareness may 549
perhaps be interpreted in a predictive coding context (Friston, 2010; Seth et al., 2011). 550
Hypothetically, high interoceptive awareness might relate to the high ‘precision’ of 552
interoceptive signals, which could, in turn, account for the high levels of autonomic 553
activity that have been observed in people with good interoceptive awareness 554
(Herbert, Pollatos, Flor, Enck, & Schandry, 2010; Pollatos, Füstös, & Critchley, 555
2012). Although very speculative, it seems possible that interoceptive signals are 556
more reliable and attended (i.e. more precise) in people with high interoceptive 557
awareness, which would account for these individuals’ reduced liability to body 558
ownership illusions. Given that interoceptive awareness affects perception of the 559
body, it is also likely to modulate action representations. It has recently been indicated 560
that in order to avoid mirroring another person’s actions it is essential to reduce the 561
precision of proprioceptive precision errors (Friston, Mattout, & Kilner, 2011). If 562
people with high interoceptive awareness have initially precise proprioceptive 563
precision errors, then their tendency to imitate others may be accounted for. 564
Potentially, recently observed individual differences in levels of neurotransmitters in 565
the insula (e.g. Wiebking et al., 2013) may provide the means to unravel the links 566
between interoceptive signals and proprioceptive, motor and autonomic reflexes. 567
5. Conclusion 569
Interoceptive awareness, measured by the accuracy with which people perceive their 570
own heartbeats, is known to modulate self/other distinction in multisensory contexts. 571
Here we demonstrate for the first time that interoceptive awareness also impacts on 572
shared representations in the motor domain, such that people with high interoceptive 573
awareness have greater difficulty in inhibiting the tendency to imitate, in a standard 574
automatic imitation paradigm. 575
6. Acknowledgements: We thank Professor Narender Ramnani for the loan of 577
equipment. 578
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... Interoception, the sense of the physiological state of the body, is thought to be critical for the subjective feeling of emotion and self-consciousness [1][2][3] . Recent studies have revealed the role of interoception in social cognition, such as sensitivity to social cues (e.g., stimuli that indicate relevant information in an interpersonal context, including others' faces, gaze, voice, or touch), imitation, and empathy in adults [4][5][6][7][8][9] , although a previous study has found no association 10 . However, little is known about the association between interoception and social cognition during infancy. ...
... Based on the hypothesis that interoception plays a role in sensitivity to social cues 13 , these findings suggest that infants' high sensitivity to interception may contribute to their perception of others' smiles as a reward, and consequently promote eye contact behavior with the mother. The present study revealed that in infants, as in adults [4][5][6][7][8][9] , interoception is associated with sensitivity to social cues, such as smiles and gaze. ...
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Interoception, that is, the perception of visceral stimuli, is the basis of socio-emotional development. However, no studies have demonstrated the relationship between the two in infants. This study aimed to elucidate the relationship between interoception and social behavior in infants and mothers. Visual preference for cardio-visual synchronous and asynchronous stimuli was assessed using a preferential-looking paradigm in 6-month infants and their mothers. The infant–mother interaction was also measured to assess social behavior, such as eye contact and positive facial expressions. The results showed that infants looked at asynchronous cardio-visual stimuli longer than synchronous cardio-visual stimuli, whereas mothers looked at synchronous cardio-visual stimuli longer than asynchronous cardio-visual stimuli. The proportion of looking time toward asynchronous cardio-visual stimuli in infants was positively correlated with infant–mother gaze and affect (positive facial expression) synchrony. Furthermore, mediation analyses showed that the relationship between infants’ interoception and eye contact behavior is attributable to mother’s positive facial expression. Our findings suggest that in infant–mother interactions, infants’ interoception may play a role in eye contact behavior through the mother's positive facial expression, highlighting the importance of infants’ interoception on social cognitive development.
... The putamen covered broader activated brain areas in the stable stage-brand love relationships than those in the other stage-brand love relationships. Given that the putamen is one of the components of the salience network (Seeley et al., 2007), connections between the anterior insula and putamen are involved in the interoceptive processing (Ainley et al., 2014;de la Fuente et al., 2019;Paik et al., 2019). Interoceptive awareness, a key element of interoceptive processing, is linked to automatic behavior (Ainley et al., 2014). ...
... Given that the putamen is one of the components of the salience network (Seeley et al., 2007), connections between the anterior insula and putamen are involved in the interoceptive processing (Ainley et al., 2014;de la Fuente et al., 2019;Paik et al., 2019). Interoceptive awareness, a key element of interoceptive processing, is linked to automatic behavior (Ainley et al., 2014). Paik et al. (2019) demonstrated that interoceptive processing plays a role in uncontrollable behavior. ...
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Brand love is a relationship between brands and consumers. Managing the relationship is an important issue for marketing strategy since it changes according to temporal flow. Brand love theories, including their dynamics, have been developed based on interpersonal romantic love theories. Although many brand love studies have provided useful findings, the neural mechanism of brand love remains unclear. Especially, its dynamics have not been considered from a neuroscience perspective. The present study addressed the commonalities and differentiations of activated brain regions between brand love and interpersonal romantic love relationships using a quantitative neuroimaging meta-analytic approach, from the view of brain connectivity. Regarding the mental processes of each love relationship related to these activated brain regions, decoding analysis was conducted using the NeuroQuery platform to prevent reverse inference. The results revealed that different neural mechanisms and mental processes were distinctively involved in the dynamics of each love relationship, although the anterior insula overlapped across all stages and the reinforcement learning system was driven between both love relationships in the early stage. Remarkably, regarding the distinctive mental processes, although prosocial aspects were involved in the mental processes of interpersonal romantic love relationships across all stages, they were not involved in the mental processes of brand love relationships. Conclusively, although common brain regions and mental processes between both love relationships were observed, neural mechanisms and mental processes in brand love relationship dynamics might be innately different from those in the interpersonal romantic love relationship dynamics. As this finding indicates essential distinctiveness between both these relationships, theories concerning interpersonal romantic love should be applied cautiously when investigating brand love relationship dynamics.
... The link between interoception and empathy has been driven by compelling meta-analytical results indicating the convergence of the neural correlates associated with interoception, emotion, and social information processing (Adolfi et al., 2017). Behavioral studies have been somewhat inconsistent, despite increasingly suggesting that interoceptive accuracy is associated with cognitive empathy (Handford et al., 2013;Ainley et al., 2014;Tajadura-Jiménez and Tsakiris, 2014;Grynberg and Pollatos, 2015;Shah et al., 2017;Shaw et al., 2020;Baiano et al., 2021). Contrastingly, evidence addressing the role of interoceptive attention within empathic processing is much scarcer and more limited (Morganti et al., 2020). ...
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This study examined the associations between psychopathy dimensions (triarchic phenotypes and classical factors), empathy domains (cognitive and affective), and interoception (interoceptive attention and accuracy) while accounting for the putative role of alexithymia. A community sample ( n = 515) completed an online survey encompassing: Triarchic Psychopathy Measure (boldness, meanness, disinhibition); Levenson Self-Report Psychopathy Scale (primary and secondary psychopathy); Body Perception Questionnaire (interoceptive attention); Interoceptive Accuracy Scale; Toronto Alexithymia Scale. Hierarchical linear regression models were implemented for hypothesis-driven analyses examining the associations between psychopathy, empathy, and interoception while controlling for sex, age, and alexithymia. Exploratory path models were employed to investigate alexithymia and/or cognitive empathy as mediators between interoception and psychopathy. Our results largely confirmed the postulated empathy profiles across psychopathy dimensions, as meanness and primary psychopathy displayed a broad empathy impairment, while disinhibition and secondary psychopathy were only associated with diminished cognitive empathy. Importantly, boldness displayed a unique pattern (enhanced cognitive empathy and reduced affective empathy), further reinforcing its importance within the constellation of psychopathy traits. Contrary to our hypotheses, self-perceived interoceptive attention and accuracy were not associated with either psychopathy dimension after controlling for alexithymia. However, interoceptive accuracy and alexithymia were associated with cognitive empathy, while alexithymia was also positively related to all psychopathy dimensions (as expected), despite the unexpected strong and negative association with boldness. Exploratory analyses suggested significant indirect effects (mediation) between interoceptive accuracy and psychopathy via alexithymia and/or cognitive empathy. These mediating effects must be interpreted with caution and future studies should be designed to formally test this model.
... Cardiac interoceptive accuracy was operationalized here as the participants' performance in the Heartbeat Counting Task (HCT) (Schandry, 1981). After a practice interval of 20 s, there were three randomized trials (25, 35, and 45 s) separated by standard resting periods of 20 s (Ainley et al., 2014). During each trial, participants were given the following instructions: "Without manually checking, can you silently count each heartbeat you feel in your body from the time you hear "start" to when you hear "stop". ...
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The influence of physical activity on interoception is apparent, however little is known about within-person variability following physical activity and sedentary behavior in daily life. To test this, 70 healthy adults (Mage 21.67±2.50) wore thigh-mounted accelerometers for 7-days, with self-reported interoception recorded on movement-triggered smartphones. Participants additionally reported the predominant activity type performed across the last 15minutes. Investigating this timeframe, multi-level analyses revealed that each one-unit increase in physical activity was associated with an increase in self-reported interoception (B = 0.0025, p =.013), whereas contrastingly, each one-minute increase in sedentary behavior was associated with a decrease (B = -0.06. p =.009). Investigating the influence of different activity types in comparison to screen time behavior, both partaking in exercise (B = 4.48, p <.001) and daily-life physical activity (B = 1.21, p <.001) were associated with an increase in self-reported interoception. Regarding other behavior categories, non-screen time behavior both with (B = 1.13, p <.001) and without (B = 0.67, p =.004) social interaction were also associated with an increase in self-reported interoception compared to screen-time behavior. Extending from previous laboratory-based studies, these findings indicate that physical activity influences interoceptive processes in real-life, further supplemented by the novel and contrasting findings regarding sedentary behavior. Furthermore, associations with activity type reveal important mechanistic information, highlighting the importance of reducing screen-time behavior to preserve and support interoceptive perceptions. Findings can be used to inform health recommendations for reducing screen-time behavior and guiding evidence-based physical activity interventions to promote interoceptive processes.
... However, mixed results have been reported regarding the relationship between interoceptive accuracy measured using the heartbeat counting task (HCT; Schandry, 1981) and the accuracy of the verbal estimation task with stimuli ranging from 19 to 103 s. Some studies have shown a moderate (Craske et al., 2001) or significant (Dunn et al., 2010;Ainley et al., 2014) correlation, while others have reported an insignificant correlation (Ehlers and Breuer, 1992;Zoellner and Craske, 1999;Dunn et al., 2007;Shah et al., 2016a,b;Murphy et al., 2018). Richter and Ibáñez (2021) demonstrated a positive correlation between interoceptive accuracy and estimated seconds of a task lasting 120 s, but not with accuracy in temporal estimation. ...
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It has been proposed that accuracy in time perception is related to interoceptive accuracy and vagal activity. However, studies investigating time perception in the supra-second range have provided mixed results, and few studies have investigated the sub-second range. Moreover, there is a lack of studies investigating the relationship between precision in time perception and interoceptive accuracy. A recent meta-analytic review of neuroimaging studies proposed a dynamic interaction between two types of timing processing—an endogenous time keeping mechanism and the use of exogenous temporal cues. Interoceptive accuracy may affect both accuracy and precision of primary temporal representations, as they are generated based on the endogenous time keeping mechanism. Temporal accuracy may vary when adapted to the environmental context. In contrast, temporal precision contains some constant noise, which may maintain the relationship with interoceptive accuracy. Based on these assumptions, we hypothesized that interoceptive accuracy would be associated with temporal precision in the sub-second range, while vagal activity would be associated with temporal accuracy. We used the temporal generalization task, which allowed us to calculate the indices of temporal accuracy and temporal precision in line with the existing research, and also compute the index of participants’ sensitivity according to the signal detection theory. Specifically, we investigated whether (1) interoceptive accuracy would correlate with temporal accuracy, temporal precision, or sensitivity and (2) resting-state vagal activity would correlate with temporal accuracy, temporal precision, or sensitivity. The results indicated that interoceptive accuracy was positively correlated with temporal precision as well as sensitivity, but not with temporal accuracy, in the sub-second range time perception. Vagal activity was negatively correlated only with sensitivity. Furthermore, we found a moderation effect of sensitivity on the relationship between vagal activity and perceived duration, which affected the association between vagal activity and temporal accuracy. These findings suggest the importance of precision as an aspect of time perception, which future studies should further explore in relation to interoception and vagal activity, and of the moderation effects of factors such as participants’ sensitivity in this context.
... Prior beliefs about resting heart rate, their actual resting heart rate, time perception ability, and BMI were measured to account for potential confounds of the Heartbeat Counting Task (HCT) [72][73][74][75] . During the time perception control task, as in previous studies 9, 70 , participants were required to estimate elapsed time duration for three randomised time intervals (23, 40, 56 s), following the same procedure as the Heartbeat Counting Task (HCT). Bivariate analyses revealed that there were no significant correlations between baseline IAcc and any of the potential confounding variables, including time perception accuracy (r = 0.226, p = 0.122), prior beliefs about HR (HR estimation accuracy) (r s = − 0.022, p = 0.884), resting HR (r = − 0.114, p = 0.441), and BMI (r = 0.227, p = 0.120). ...
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Interoception, the sense of the internal body, is proposed to support self-regulation, and consequently influence mental health. Researchers have therefore shown interest in improving the ability to accurately monitor internal signals (i.e., interoceptive accuracy, IAcc). Research suggests that cardiac IAcc is modifiable by both manipulations of interoceptive attention (guided attention towards the internal body), and interoceptive exposure (strategically inducing somatic signals e.g., via physical activity). Whilst successful in isolation, it is unclear whether a combined approach (i.e., directing attention towards the internal body when signals are more salient) could elicit greater benefits. In a 2 × 2 within-subject design, 48 healthy adults ( M age = 25.98 ± 4.73 years, 50% female) completed four 20-min conditions varying in both attentional focus (interoceptive vs exteroceptive) and physical activity (active vs rest), with cardiac IAcc measured immediately after. Results revealed a main effect for physical activity ( p < 0.001), however, there was no effect for attentional focus ( p = 0.397), and no interaction effect ( p = 0.797). Differential analyses showed that a higher sporting background increased sensitivity to physical activity-related increases in cardiac IAcc ( p = 0.031). Findings indicate that (irrespective of attentional focus) moderate-vigorous physical activity-based interventions have the potential to increase cardiac IAcc, with certain individuals potentially benefiting more.
... Additionally, heartbeats may be perceived via vibrations against the chest wall (Khalsa et al., 2009) and depend on factors such as body-fat percentage (Rouse et al., 1988), and resting heart-rate variability (HRV; Knapp-Kline & Kline, 2005). Some questioning the validity and reliability of the HCT (Brener & Ring, 2016;Desmedt et al., 2018) also suggested that participants may report their felt heartbeats based on an approximation of the time they think has passed, general knowledge about average heartbeats in the general population and a calculation of the number of heartbeats they should have felt based on time awareness and heartbeat knowledge (e.g., Ainley et al., 2014;Murphy, Geary et al., 2018;Shah et al., 2016). Though, Murphy et al. (2018b) suggested that the criticisms may be less valid when controlling for these aforementioned measuresmeasures which we will control for in our follow-up study. ...
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Anorexia Nervosa (AN) is an eating disorder with high mortality and morbidity rates, partly due to treatment resistance and high relapse rates. Treatment adherence and recovery has been found to be hindered by insight deficits, a lack of appreciation of one’s illness, or its consequences, most frequent in restrictive AN. However, to date, insight disturbances in AN have mainly been studied in relation to treatment outcomes rather than explanatory mechanisms. One possibility is that interoception (the sensing, awareness and interpretation of physiological signals) and particularly its metacognitive aspects such as prospective (self-efficacy) and retrospective (insight) beliefs about one’s interoceptive abilities may be affected in AN. To our knowledge however such aspects of global metacognition, and their relation to key interoceptive and body perception impairments, have not been assessed in AN. Here in two experiments (nAN=51 and 28, nAN-WR=47 and 21, nHC=63 and 34, respectively), we tested, (a) how women with and weight-restored from AN (AN-WR), in comparison to healthy controls (HCs), formulate explicit interoceptive self-efficacy beliefs (i.e., estimates of performance in a cardiac perception task) prospectively and then update them following performance and then following explicit feedback and (b) how they formulate prospectively and then update following feedback two types of body-size beliefs (estimates about the envisioned body, ‘How thin it looks' vs the emotional body, ‘How thin it feels’). Results of Experiment 1 confirmed our hypotheses that the AN (but not the AN-WR) group formulated more pessimistic interoceptive self-efficacy beliefs in comparison to HCs both before and after otherwise comparable performance. In Experiment 2 we found that the AN group envisioned and felt (also the AN-WR group) their body size to be bigger than it really is in comparison to controls. Post-feedback, the AN but not AN-WR group significantly overestimated both their envisioned and emotional body and they also updated their emotional body size estimates at a slower rate than the HCs. These observed group differences in belief updating about interoceptive self-efficacy and body size estimates warrant further studies in interoceptive metacognition and belief updating in AN, and their relation with insight deficits, particularly at the acute stages of the disease.
... Note that this a priori power analysis was based on a simpler paradigm (four conditions in 2 × 2 factorial with the same stimulus-response action pairs), so to accommodate for the additional complexity of the current paradigm and analysis plan, including trial-wise modelling, we targeted a sample of 30 participants. This sample also aligns with the sample size used in other multifaceted SRC tasks (e.g., Ainley et al., 2014). ...
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Dyadic interactions require dynamic correspondence between one’s own movements and those of the other agent. This mapping is largely viewed as imitative, with the behavioural hallmark being a reaction-time cost for mismatched actions. Yet the complex motor patterns humans enact together extend beyond direct-matching, varying adaptively between imitation, complementary movements, and counter-imitation. Optimal behaviour requires an agent to predict not only what is likely to be observed but also how that observed action will relate to their own motor planning. In 28 healthy adults, we examined imitation and counter-imitation in a task that varied the likelihood of stimulus–response congruence from highly predictable, to moderately predictable, to unpredictable. To gain mechanistic insights into the statistical learning of stimulus–response compatibility, we compared two computational models of behaviour: (1) a classic fixed learning-rate model (Rescorla–Wagner reinforcement [RW]) and (2) a hierarchical model of perceptual-behavioural processes in which the learning rate adapts to the inferred environmental volatility (hierarchical Gaussian filter [HGF]). Though more complex and hence penalized by model selection, the HGF provided a more likely model of the participants’ behaviour. Matching motor responses were only primed (faster) in the most experimentally volatile context. This bias was reversed so that mismatched actions were primed when beliefs about volatility were lower. Inferential statistics indicated that matching responses were only primed in unpredictable contexts when stimuli–response congruence was at 50:50 chance. Outside of these unpredictable blocks the classic stimulus–response compatibility effect was reversed: Incongruent responses were faster than congruent ones. We show that hierarchical Bayesian learning of environmental statistics may underlie response priming during dyadic interactions.
L’autore esplora il moderno concetto di interocezione ed il ruolo delle atipie interocettive quale fattore crossdiagnostico a differenti disturbi mentali, al fine di presentarne le implicazioni per la psicoterapia ericksoniana. Vengono esaminati i processi interocettivi, in termini di dimensioni, caratteristiche e traiettorie di sviluppo, evidenziandone il ruolo centrale nell’esperienza emozionale. Viene presentato come osservare, valorizzare e utilizzare le modificazioni dei processi interocettivi in ipnosi ed esemplificato, attraverso casi clini-ci, come tale plasticità possa promuovere una rielaborazione globale dell’esperienza emozionale nel contesto del rapport.
People tend to perceive a virtual body standing in front of them as their own if it is either stroked or moving synchronously with their own real body—the out-of-body experience (OBE). We combined synchrony manipulation with two other factors of theoretical interest: the kind of stimulation, visuotactile stimuli or visuomotor correlations, being synchronized and the interoceptive accuracy (IA) of participants, assessed by means of the heartbeat-detection task. Results showed that explicit measures of embodiment were systematically affected by synchrony, and this synchrony effect was more pronounced for visuomotor than for visuotactile. The walking drift perception was affected by IA: In visuotactile conditions, the synchrony effect was pronounced in individuals with low IA, presumably reflecting a stronger impact of the visual information. In visuomotor, however, the synchrony effect was stronger in individuals with high IA, presumably reflecting a stronger impact of re-afferent information generated by the participants’ own movements.
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The individual sensitivity for ones internal bodily signals ("interoceptive awareness") has been shown to be of relevance for a broad range of cognitive and affective functions. Interoceptive awareness has been primarily assessed via measuring the sensitivity for ones cardiac signals ("cardiac awareness") which can be non-invasively measured by heartbeat perception tasks. It is an open question whether cardiac awareness is related to the sensitivity for other bodily, visceral functions. This study investigated the relationship between cardiac awareness and the sensitivity for gastric functions in healthy female persons by using non-invasive methods. Heartbeat perception as a measure for cardiac awareness was assessed by a heartbeat tracking task and gastric sensitivity was assessed by a water load test. Gastric myoelectrical activity was measured by electrogastrography (EGG) and subjective feelings of fullness, valence, arousal and nausea were assessed. The results show that cardiac awareness was inversely correlated with ingested water volume and with normogastric activity after water load. However, persons with good and poor cardiac awareness did not differ in their subjective ratings of fullness, nausea and affective feelings after drinking. This suggests that good heartbeat perceivers ingested less water because they subjectively felt more intense signals of fullness during this lower amount of water intake compared to poor heartbeat perceivers who ingested more water until feeling the same signs of fullness. These findings demonstrate that cardiac awareness is related to greater sensitivity for gastric functions, suggesting that there is a general sensitivity for interoceptive processes across the gastric and cardiac modality.
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A surfeit of research confirms that people activate personal, affective, and conceptual representations when perceiving the states of others. However, researchers continue to debate the role of self-other overlap in empathy due to a failure to dissociate neural overlap, subjective resonance, and personal distress. A perception-action view posits that neural-level overlap is necessary during early processing for all social understanding, but need not be conscious or aversive. This neural overlap can subsequently produce a variety of states depending on the context and degree of common experience and emotionality. We outline a framework for understanding the interrelationship between neural and subjective overlap, and among empathic states, through a dynamic-systems view of how information is processed in the brain and body.
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Previous research has yielded inconsistent results concerning affective reactions to social exclusion. The present study provides evidence that conscious perception of bodily signals (“interoceptive awareness”) constitutes an important moderating factor in this context. We compared participants with high versus low cardiac interoceptive awareness in regard to affective, cognitive, and physiological measures while they were included and excluded in a discussion with confederates. Participants with high interoceptive awareness showed a smaller decrease of positive affect and perceived acceptance as well as a smaller increase of negative affect and perceived rejection when comparing an inclusion phase with a subsequent exclusion phase than did participants with low interoceptive awareness. No significant differences in cognitive and physiological measures were observed. We assume that individuals with high interoceptive awareness, to whom physiological signals are more easily accessible, reduce aversive states to a larger degree by using somatic information for self-regulation. (PsycINFO Database Record (c) 2013 APA, all rights reserved)
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We spend much of our life predicting the future. This involves developing theories and making predictions about others' intentions, goals and about the consequences of the actions we are observing. Adapting our actions and behaviours to the environment is required for achieving our goals, and to do this the motor system relies on input from sensory modalities. However, recent theories suggest that the link between motor and perceptual areas is bidirectional, and that predictions based on planned or intended actions can unconsciously influence and modify our perception. In the following review we describe current theories on the link between action and perception, and examine the ways in which the motor system can unconsciously alter our perception.
To facilitate a multidimensional approach to empathy the Interpersonal Reactivity Index (IRI) includes 4 subscales: Perspective-Taking (PT) Fantasy (FS) Empathic Concern (EC) and Personal Distress (PD). The aim of the present study was to establish the convergent and discriminant validity of these 4 subscales. Hypothesized relationships among the IRI subscales between the subscales and measures of other psychological constructs (social functioning self-esteem emotionality and sensitivity to others) and between the subscales and extant empathy measures were examined. Study subjects included 677 male and 667 female students enrolled in undergraduate psychology classes at the University of Texas. The IRI scales not only exhibited the predicted relationships among themselves but also were related in the expected manner to other measures. Higher PT scores were consistently associated with better social functioning and higher self-esteem; in contrast Fantasy scores were unrelated to these 2 characteristics. High EC scores were positively associated with shyness and anxiety but negatively linked to egotism. The most substantial relationships in the study involved the PD scale. PD scores were strongly linked with low self-esteem and poor interpersonal functioning as well as a constellation of vulnerability uncertainty and fearfulness. These findings support a multidimensional approach to empathy by providing evidence that the 4 qualities tapped by the IRI are indeed separate constructs each related in specific ways to other psychological measures.
To remove the influence of spuriously long response times, many investigators compute "restricted means," obtained by throwing out any response time more than 2.0, 2.5, or 3.0 standard deviations from the overall sample average. Because reaction time distributions are skewed, however, the computation of restricted means introduces a bias: the restricted mean underestimates the true average of the population of response times. This problem may be very serious when investigators compare restricted means across conditions with different numbers of observations, because the bias increases with sample size. Simulations show that there is substantial differential bias when comparing conditions with fewer than 10 observations against conditions with more than 20. With strongly skewed distributions and a cutoff of 3.0 standard deviations, differential bias can influence comparisons of conditions with even more observations.
Assessed heart beat perception across diagnostic subgroups of psychosomatic disorders. 546 inpatients (mean age 40.9 yrs), distributed into diagnostic subgroups (neurotic depressives, depressive reactions, functional disorders, personality disorders, functional cardiac disorders, and panic disorders), were compared to 48 health controls (mean age 35.8 yrs) in heart beat perception over an interval of 4 wks. Medication effects and sex differences were also examined. Marginally significant differences in heart beat perception scores between subgroups were evident. The detection score of patients with personality disorders was lower compared to patients with functional disorders and control Ss. Functional heart disorder and panic patients did not have higher perception scores than control Ss. Medicated panic patients showed better heart beat perception than unmedicated ones. The distribution of heart beat perception was similar in all groups studied. In general, psychosomatic patients tended toward lower perception scores compared to control Ss. Findings suggest that better than normal interceptive functioning does not seem to play a decisive role in the establishment of functional psychosomatic disorders. (PsycINFO Database Record (c) 2012 APA, all rights reserved)
Mark Solms suggests that the internal body, equated with the id and represented at the brainstem structures, gives us the self-as-subject consciousness, while the external body, equated with the ego, is represented as an object, analogous to any other object in the world. Is memory space the sole, or at least the most important, contribution that the ego can make to the id? I would like to argue that it is not. Even though the basis of phenomenal consciousness—the “being-me” state—might be given by the brainstem consciousness, the most important function of the ego is precisely that it can represent my body as an object and identify it with the internal body. Thus, both bodies need to be represented as self-specific, and inevitably this will require the contribution of a cortical network. Both the ego and the id, in Solms’s terms, co-constitute self-specificity.