Inhibitory control training might be a gateway to enhance fear extinction.

Inhibitory learning forms an essential component of extinction learning. Deficits in inhibitory learning could negatively impact extinction learning. Previous research has shown optimizing inhibitory learning has potential to improve extinction. The current study aims to improve inhibitory learning capacity through training inhibitory control to improve extinction. Our results show that training inhibitory control through stop-signal task improves extinction learning as evidenced through better reduction in UCS-expectancy, further it also resulted in reduced subjective arousal ratings during extinction. This effect was persistent and observed 24 hours later during extinction recall. Our findings imply that training inhibitory control assists in forming inhibitory associations during extinction. Further, we also observed elevated physiological arousal in training group during extinction, this explains the better recall of extinction learning during extinction recall phase. Previous research has shown better consolidation of memory under high arousal state. Improving individual capacity to form inhibitory associations enhances extinction, this finding could aid developing interventions for individuals with anxiety disorders, who show deficits in inhibitory learning. Additionally, our results also provide support to retrieval stopping hypothesis of extinction, and highlighting the domain general nature of inhibitory control involved in motoric inhibition and inhibitory processes involved in extinction. Inhibitory control training might be a gateway to enhance fear extinction. Kaneez Fatima Dar 1 and Manish Kumar Asthana 1,2 * 1 Department of Humanities & Social Sciences, Indian Institute of Technology Roorkee, India 2 Department of Design, Indian Institute of Technology Roorkee, India * Corresponding author ([email protected], [email protected]) First Author ([email protected]) Declaration of conflicting interest The authors declare that they have no conflict of interest.

Inhibitory learning forms an essential component of extinction learning. Deficits in inhibitory learning could negatively impact extinction learning. Previous research has shown optimizing inhibitory learning has potential to improve extinction. The current study aims to improve inhibitory learning capacity through training inhibitory control to improve extinction. Our results show that training inhibitory control through stop-signal task improves extinction learning as evidenced through better reduction in UCS-expectancy, further it also resulted in reduced subjective arousal ratings during extinction. This effect was persistent and observed 24 hours later during extinction recall. Our findings imply that training inhibitory control assists in forming inhibitory associations during extinction. Further, we also observed elevated physiological arousal in training group during extinction, this explains the better recall of extinction learning during extinction recall phase. Previous research has shown better consolidation of memory under high arousal state. Improving individual capacity to form inhibitory associations enhances extinction, this finding could aid developing interventions for individuals with anxiety disorders, who show deficits in inhibitory learning. Additionally, our results also provide support to retrieval stopping hypothesis of extinction, and highlighting the domain general nature of inhibitory control involved in motoric inhibition and inhibitory processes involved in extinction.

fear conditioning, inhibitory control, inhibitory learning, fear extinction, stop-signal task

Extinction is a process aimed at reducing conditioned fear responses through repeated exposure to feared stimuli without the aversive outcome, signalling that it is no longer something to fear (Bouton et al., 2006). Inhibitory learning forms an essential process of extinction learning (Bouton, 1993), although other mechanism like habituation is also proposed to be involved (Myers & Davis, 2007). Based on habituation account of extinction, decrease in conditioned responses during extinction results from repeated exposure to conditioned stimulus (McSweeney & Swindell, 2002). Inhibitory learning involves learning that feared stimuli are not always followed by an aversive outcome. Extinction after repeated exposure results in lower fear responses and together with inhibitory learning individuals learn that a stimulus might not always signal threat, resulting in lower fear responses in response to the stimulus encountered in a new context (Lebois et al., 2019). Incidences of emergence of fear learning after extinction through spontaneous recovery (re-exposure to CS+ after certain time), US re-exposure (UCS presentations) and renewal (change in context), suggests extinction learning does not result from the complete erasure of CS+-UCS association but rather results from the formation of an inhibitory association (Bouton, 2002). After extinction, the original CS-UCS association formed after fear conditioning remains intact, however a secondary CS-no UCS association is formed (Craske et al.2014). After inhibitory learning during extinction the new extinction memory is stored through the process of consolidation (Tronson & Taylor 2007). Previous research suggests that low inhibitory capacity might delay extinction, due to slower formation of new inhibitory associations (CS+- not followed by UCS) resulting in expression of excitatory association (CS+ followed by UCS). Excitatory associations result in automatic fear responses which are difficult to inhibit (Miyake, 2000). This is corroborated by previous work suggesting that highly anxious individuals (Lissek et al., 2005) fail to repress physiological and self-reported fear to stimuli. Anxiety disorders are associated with heightened excitatory processes and impaired inhibitory learning processes. Individual experience relapse in fear or inadequate symptom relief following exposure therapy (Arch & Craske, 2009). This in part is attributed to inadequate extinction learning particularly inhibitory learning and associated inhibitory neural regulation during extinction. Therefore, improving inhibitory learning has the potential to enhance extinction outcomes (Craske et al., 2014). Previous studies dealing with improving extinction through inhibitory learning mechanism have engaged strategies like expectancy violation (Baker et al., 2010), deepened extinction (Rescorla, 2006), occasional reinforced extinction (Bouton et al., 2004) etc. The focus of the current study is on individual capacity to form negative associations and improving such skills through training. Inhibitory control is a component of executive functions and is understood as an individual capacity to restrain behavioural impulses and affective states (Venables et al., 2018). Inhibitory control makes sure we are not vulnerable to our impulses or old ways of thought or action (conditioned responses) (Diamond, 2014). Inhibitory phenomenon in associative learning is distinguishable at various levels and contexts: inhibitory learning, response inhibition and inhibitory control. Inhibitory learning involves changes in behaviour resulting from learning negative associations between events. Response inhibition is implementation of learning of these negative association and inhibitory control is defined as an individual trait expressing the capacity of forming such negative association (Sosa et al., 2022). Since these concepts are interlinked, we hypothesized that training on response inhibition task would improve inhibitory control which consequently would improve inhibitory learning. Inhibitory learning is central to extinction, improving individual capacity to regulate inhibition could help in improve formation of inhibitory association resulting in better extinction learning. Previous studies have revealed that high resting heart rate variability, an indicator of autonomic inhibitory control is associated with better extinction (Pappens et al., 2014). This has been extended to generalization phenomenon as well, showing that impaired inhibitory capacity results in slower extinction of generalization stimuli (Niederstrasser et al., 2017). In our study we investigated the effect of training inhibitory control through stop-signal task, a common behavioural measure of inhibitory control (Houben, Nederkoorn, & Jansen, 2014; Liu, Roefs & Nederko0rn, 2022). We hypothesize that as inhibitory control strengthens, it may enhance the individual’s capacity to recognize and learn from negative associations more effectively. Further, according to retrieval stopping hypothesis of extinction (Anderson & Floresco, 2022), extinction involves cognitive control processes, it recruits retrieval stopping processes. Retrieval stopping is similar to action stopping in terms that both engage right dorsolateral and ventrolateral prefrontal cortexes. Retrieval stopping involves prefrontal cortex mediated suppression of amygdala and hippocampus, whereas action stopping involves suppression of motor cortex. This similar recruitment of lateral prefrontal cortex in both retrieval and action stopping suggests a domain-general mechanism involved in inhibition of varied content through shift in connectivity to areas requiring control. It also hypothesizes that extinction involves activation of right dorsolateral prefrontal cortex similar to retrieval stopping, although extinction has been mostly associated with vmPFC activation. Fullana et al., in their meta-analysis observed activation of rDLPC during extinction. Based on the assumption that extinction involves retrieval stopping, it could be hypothesized that improving or training inhibitory control in one context (action stopping) could result in improvement in other (extinction). Methodology Sample size Sample size was determined using a priori power analysis with an alpha value of 0.05, power of 0.80, and a medium effect size of 0.25, the sample size required was 30 using repeated measures ANOVA for measures recorded after each phase. Participants 52 healthy participants (21 females, 31 males; M= 23.07, SD=0.56) completed the three-day experiment. One participant was excluded from analysis due to no measurable SCR recorded on day 1, the analysis included 51 participants (21 females, 31 males; M= 23.16, SD=4.06). Exclusion criteria for the study were as follows: people currently using any medication that can impair attention, concentration, reaction time, or memory; diagnosed with any psychiatric disorder like anxiety, phobia, depression, etc.; under the treatment of any mental health professional in the past two years; pregnant women ; diagnosed with any serious neurological or medical condition like epilepsy or heart disease; have consumed alcohol, any psychoactive substances or more than 5mg of caffeinated drinks in the past 24 hours. The project was approved by the Institute Human Research Ethics Committee, Indian Institute of Technology Roorkee (IITR/ICC/24/18). Experimental design Before the experiment began participants were asked for written consent for their participation in the experiment. Participants later filled out two questionnaires consisting of State-trait Anxiety Inventory and Emotion regulation questionnaire. We used the Screaming Lady paradigm (Lau et al., 2009). A three-day differential fear conditioning paradigm was used for the experiment. Day 1 comprised of habituation and acquisition, day 2 of extinction and day 3 of extinction recall. Two neutral black and white female faces from NimStim database served as CS+ and CS- (Lau et al., 2008; Ney, J. et al.,2021). Habituation and Acquisition occurred on day 1 with 4 trials of each CS in habituation and 16 trials of each CS in acquisition on, CS+ on 75% of the trials was followed by a UCS (fearful expression face and scream) for 3 seconds. Extinction consisted of 12 trials of each CS, without CS+ being followed by UCS. Extinction recall took place on with 6 trials of each CS. The trials in each phase were presented pseudorandomly and the CS+ stimuli was counterbalanced across participants. Each CS was presented for 8 seconds, followed by a 10 to 12 seconds inter-trial interval (Figure 1). The experiment consisted of two groups: Inhibitory control training and extinction group (IC-SE) and extinction only group. All the groups followed the same procedure except for day 2, in inhibitory control training and extinction group (IC-SE) participants were trained on stop-signal task followed by extinction. Whereas, the extinction only group on day 2 didn’t undergo any inhibitory control training. Fear conditioning stimuli Conditioned stimuli (CSs) Two neutral and black and white female faces from NimStim (03F_NE_C, 18F_NE_C) database were used as either CS+ and CS- (Lau et al., 2008; Ney, J. et al.,2021). The CS+ and CS- neutral images were presented for 8 seconds and assignment of face stimuli as CS+ and CS- was counterbalanced. The presentation of CS+ and CS- was pseudorandomized with a randomized intertrial interval (ITI) of 10 to 12 seconds. Unconditioned stimulus (UCS) The UCS was a fearful expression image of the same model (03F_FE_O, 18F_FE_O) as CS+ paired with a 95-dB scream (Femscream3, Sound no.: 277) from Internation affective digitized sounds (IADS-2) (Bradley & Lang, 2007), presented for 3 seconds (Lau et al., 2008; Culver et al; 2018; Heather & Waters, 2019). The screaming sound was delivered through headsets. Figure 1 Schematic of experimental phases (A) Acquisition, (B) Extinction and Extinction recall Inhibitory control training As an inhibitory control training, we used a response inhibition task. We opted for a response inhibition task since its results in inhibition of action that are no longer relevant (Verbruggen & logan, 2009). The stop-signal task consisted of a total 240 trials with 180 go-trials which formed 75% of total trials and 60 stop -trial which was a total of 25% of trials (Verbruggen et al., 2019). Before the training block participants were presented with a practice block consisting of 24 trials with 6 stop signal trials. The training took place in 6 blocks, participants were allowed to take break before the beginning of each block. Each block consisted of 40 trials with 10 stop trials in each block. In Go-trials, participants were requested to respond as quickly and accurately as possible to the direction of the black arrow appearing on the screen for 1500 ms. Each arrow was followed by a fixation cross of 500 ms duration (Ding et al., 2020). The participants were instructed to respond with ‘p’ when the direction of the arrow was towards right and to respond with ‘q’ if the direction of the arrow was towards left using the keyboard. The presentation of the stop-trials and go-trials was randomized. During stop-trials, participants would hear beep meaning that they have to inhibit their motor response after a variable stop-signal delay (SSD) relative to the onset of the Go-stimulus. In the beginning of the task, the beep was emitted 250 ms following the display of the arrow. This delay increased by 50 ms when the response was successfully inhibited and decreased by 50 ms when the response was not inhibited. The outcome variable for this task was the stop signal reaction time (SSRT). The latter was computed by subtracting averaged stop signal delay from the mean reaction time on go trials. A longer SSRT is indicative of lower response inhibitory capacity (Niederstrasser et al., 2016) (Figure 2). Figure 2 Schematic of stop-signal task (A) Go-trials (b) Stop-trials Subjective measures Trait Anxiety measure Before the experiment the participants filled out the State-Trait Anxiety Inventory for Adults (STAI; Spielberger, Gorsuch, Lushene, Vagg, & Jacobs, 1983). This was used to assess anxiety symptoms. The STAI is a self-report scale which comprises of two 20-item scales designed to assess ”state anxiety” which is referred to as temporary experience of anxiety symptoms and ”trait anxiety” which indicates consistent anxiety symptoms. Each subscale score ranges from 20 to 80, with higher scores indicating higher anxiety symptoms. The participants filled out STAI trait anxiety questionnaire. CS valence and arousal Participants were asked to rate the valence and arousal of both CS at the end of phase on a 9-point rating scale with self-assessment mannequin figures (Bradley & Lang, 1994; O’Malley & Waters, 2018). The valence scale ranged from 1(unpleasant) to 9 (pleasant) and arousal scale ranged from 1(calm) to 9 (excited). The scale was presented under an image of CS, and participants were instructed to record their response by pressing the appropriate number on the keyboard. Subjective fear scale Participants rated the subjective fear to both CS at the end of each phase using a 9-point rating scale. The fear scale ranged from 1(low) to 5 (moderate) to 9 (high) (Hermann, Keck & Stark, 2014). The scale was presented under the CS image and participants were instructed to record their response by pressing the appropriate number on the keyboard. Expectancy scale To indicate the UCS expectancy participants were presented with a 9-point rating scale (1= Highly unlikely, 5= Uncertain, 9 = Highly likely) for each CS presentation (Constantinou et al., 2021). Participants were asked to responds using the keyboard. Skin conductance response The physiological responses were measured as skin conductance response (SCR) recorded through Biopac MP160 (BIOPAC Systems, Inc., Goleta, California, USA) and data was analysed using Acknowledge 3.9 software. SCR was acquired through two Ag-Agcl reusable electrodes attached to the distal phalanges of the second and third fingers of participant’s non-dominant hand. SCR data was filtered using Finite Impulse Response (FIR) low pass filter at frequency cut-off of 1Hz. The data was segregated into different focus areas according to experimental phases (habituation, acquisition, extinction and extinction recall). The digital inputs were converted to stimulus events and a phasic EDA channel was set up using 0.05 Hz high pass filter. SCR was computed as base-to-peak amplitude in 1 to 8s window following the CS onset. The SCR minimal response criterion was set at 0.02 μS responses below this criterion were recorded as zero (Shiller et al., 2013, Woelk et al., 2021). The raw SCR data was square root transformed and range corrected (Boucsein et al., 2012).

Table 1 Descriptive data of trait anxiety scale | STAI-Trait (M, (SD)) | 41.4 (7.1) | 44.5(9.4) | .185 | -.337 | Table 2 Descriptive data from stop-signal Analysis The statistical analysis of data was conducted using IBM SPSS software version 27. For analysis the first trial of CS- was removed from each phase as orientation response for each measurement. We conducted a t-test to compare the trait anxiety of participants between the groups, the results revealed no significant difference on trait anxiety between the groups (Table 1). The stop-signal reaction time (SSRT) was computed by subtracting averaged stop signal delay from the mean reaction time on go trials (Table 2). Arousal ratings We performed a 4 x 2 x 2 repeated measure ANOVA with phase (habituation, acquisition, extinction and extinction recall) and stimulus type (CS+. CS-) as within subject factor and group (IC-SE & SE) as between subject factor on arousal ratings. The results indicated a significant main effect of phase [F (2.45,120.16) = 17.28, p< .001, ƞp2 =.261] 1, further, a significant main effect of stimulus type [F (1,49) = 28.76, p< .001, ƞp2 =.370]. Additionally, the results also indicated a significant phase x stimulus type interaction [F (2.29,112.12) = 15.35, p< .001, ƞp2 =.239; see Footnote 1], and a significant phase x stimulus type x group interaction [F (2.29,112.12) = 3.92, p< .018, ƞp2 = .074; see Footnote 1]. Further, a pairwise comparison of phase revealed a significant difference between habituation (M= 3.42) and acquisition (M= 4.50) phase (p< .001, 95% CI (-1.64, -.52), further, acquisition (M= 4.50) differed significantly from extinction (M= 3.49) (p< .001, 95% CI (.43, 1.60) and extinction recall phase (M= 3.00) (p< .001, 95% CI (.78, 2.22). Extinction (M= 3.49) differed significantly from extinction recall (M= 3.00) phase (p = .027, 95% CI (.04, .96). Additionally, a pairwise comparison of stimulus revealed a difference in CS+ (M= 4.32) and CS- (M= 2.89) (p< .001, 95% CI (.89, 1.96)). To explore the interaction effect further we performed a 2 x 2 repeated measures ANOVA with stimulus type (CS+, CS-) as within subject factor and group (IC-SE, SE) was conducted on arousal ratings for each phase separately. In habituation phase the results indicated non-significant main effect of stimulus type [F (1,49) = .047, p= .829, ƞp 2 =.001] indicating no difference in baseline arousal in CS+ (M=3.392, SD= 1.990) and CS- (M=3.431, SD= 2.022) during habituation. Further, no significant stimulus x group interaction effect was observed. In acquisition phase the results indicated a significant main effect of stimulus type [F (1,49) = 23.977, p < .001, ƞp 2 =.329] with higher arousal for CS+ (M=5.666, SD= 2.581) than CS- (M=3.294, SD= 2.220) this indicates successfully fear learning. Further, no significant stimulus x group interaction effect was observed, implying comparative fear acquisition in both groups. In extinction phase the results indicated a significant main effect of stimulus type [F (1,49) = 31.539, p < .001, ƞp 2 =.392] with higher arousal for CS+ (M= 4.392, SD= 2.010) than CS- (M= 2.549, SD= 1.921). Further, a significant stimulus type x group interaction effect was observed [F (1,49) = 4.832, p = .033, ƞp 2 =.090]. An independent sample t-test was conducted to assess the group difference on CS+ and CS- arousal ratings, the results revealed a significant group difference on CS+ arousal rating t (49) = -2.760, p = .008, d = -.774 with lower CS+ arousal ratings in IC-SE group (M =3.703, SD= 1.835) than SE group (M= 5.166, SD = 1.948). There was no difference on CS- arousal ratings between the groups. Further, an independent sample t-test also revealed a significant group difference in differential arousal ratings, calculated as CS+-CS- arousal values t (49) = -2.198, p = .033, d = -.617, with lower differential value in IC-SE group (M= 1.148, SD = 2.298) than SE group (M= 2.625, SD = 2.498). In extinction recall phase the results indicated a significant main effect of stimulus type [F (1,49) = 35.818, p < .001, ƞp 2 =.422] with higher arousal for CS+ (M= 3.705, SD= 1.746) than CS- (M= 2.254, SD= 1.647). Further, a significant stimulus x group interaction effect was observed [F (1,49) = 6.552, p = .014, ƞp 2 = .118]. An independent sample t-test was conducted to assess the group difference on CS+ arousal ratings, the results revealed a significant group difference on CS+ arousal rating t (49) = -2.548, p = .014, d= -.715 with lower CS+ arousal ratings in IC-SE group (M = 3.148, SD= 1.633) than SE group (M= 4.333, SD = 1.685). Further, an independent sample t-test also revealed a significant group difference in differential arousal ratings t (49) = -2.560, p = .014, d= -.718, with lower differential value in IC-SE group (M= .851, SD = 1.854) than SE group (M= 2.125, SD = 1.676) (Figure 3). Figure 3 Subjective arousal plots by (A) differential values (B) CS Type Expectancy ratings We performed a 3 x 2 x 2 repeated measure ANOVA with phase (acquisition, extinction and extinction recall) and stimulus type (CS+, CS-) as within subject factor and group (IC-SE & SE) as between subject factor on UCS-expectancy ratings. The results indicated a significant main effect of phase [F (1.73,86.34) = 91.208, p< .001, ƞp2 =.646; see Footnote 1], and a significant main effect of stimulus type [F (1,50) = 232.81, p< .001, ƞp2 =.823]. Additionally, the results also indicated a significant stimulus type x group interaction [F (1, 50) = 5.143, p= .028, ƞp2= .093] and significant phase x stimulus type interaction [F (1.60,79.96) = 21.96, p< .001, ƞp2 =.305; see Footnote 1]. Further, a pairwise comparison of phase revealed that acquisition phase (M= 5.28) differed significantly from extinction (M= 3.73) (p< .001, 95% CI (1.140, 1.951) and extinction recall (M= 3.014) phase (p< .001, 95% CI (1.765, 2.761). Extinction (M= 3.73) differed significantly from extinction recall (M= 3.014) phase (p< .001, 95% CI (.361, 1.075). Additionally, a pairwise comparison of stimulus revealed a difference in CS+ (M= 5.677) and CS- (M= 2.338) (p< .001, 95% CI (2.899, 3.778)). To further explore the interaction effect, we performed a 2 x 2 repeated measures ANOVA with stimulus type (CS+, CS-) as within subject factor and group (IC-SE, SE) on UCS- expectancy ratings for each phase separately. In acquisition phase the results indicated a significant main effect of stimulus type [F (1,49) 224.984=, p < .001, ƞp 2 =.821] with higher UCS expectancy for CS+ (M= 7.438, SD= 1.233) than CS- (M= 3.099, SD= 1.541) this indicates successfully fear learning. Further, no significant stimulus x group interaction effect was observed, implying comparative fear acquisition in both groups. In extinction phase the results indicated a significant main effect of stimulus type [F (1,49) =119.410, p < .001, ƞp 2 =.709] with higher UCS expectancy for CS+ (M= 5.261, SD=2.126) than CS- (M= 2.163, SD= 1.428). Further, a significant stimulus x group interaction effect was observed [F (1,49) = 4.703, p = .035, ƞp 2 =.088]. An independent sample t-test was conducted to assess the group difference on CS+ and CS- expectancy ratings, the results revealed a significant group difference on CS+ expectancy rating t (49) = -2.211, p = .032, d = -.620, with lower CS+ expectancy ratings in IC-SE group (M = 4.663, SD= 2.045) than SE group (M= 5.934, SD =2.051). The results revealed a non-significant group difference on CS- expectancy values. In extinction recall phase the results indicated a significant main effect of stimulus type [F (1,49) = 86.851, p < .001, ƞp 2 =.639] with higher UCS expectancy for CS+ (M= 4.241, SD= 2.145) than CS- (M= 1.803, SD= 1.344). Further, a significant stimulus x group interaction effect was observed [F (1,49) = 5.684, p = .021, ƞp 2 = .104]. An independent sample t-test was conducted to assess the group difference on CS+ and CS- expectancy ratings, the results revealed a significant group difference on CS+ expectancy rating t (49) = -2.567, p = .011, d= -.746, with lower CS+ expectancy ratings in IC-SE group (M = 3.530, SD= 1.886) than SE group (M= 5.041, SD = 2.173). The results revealed a no significant group difference on CS- expectancy values (Figure 4). Figure 4 UCS-expectancy plots by CS-type (A) acquisition, (B) extinction and (C) extinction recall phase Skin Conductance response values We performed a 4 x 2 x 2 repeated measure ANOVA with phase (habituation, acquisition, extinction and extinction recall) and stimulus type (CS+. CS-) as within subject factor and group (IC-SE & SE) as between subject factor on SCR ratings. The results indicated a significant main effect of phase [F (2.44, 119.58) = 4.512, p = .008, ƞp2 = .084; see Footnote 1], further, the results also indicated a non- significant main effect of stimulus type [F (1,49) = 2.473, p= .122, ƞp2 = .048]. Additionally, the results indicated a significant phase x stimulus type interaction [F (2.56,126.17) = 5.012, p = .004, ƞp2 = .093; see Footnote 1], additionally a non- significant phase x stimulus type x group was found [F (2.56,126.17) = .088, p =.951, ƞp2 = .002; see Footnote 1]. Further, a pairwise comparison of phase revealed a significant difference between habituation (M= .129) and acquisition (M= .174) phase (p< .017, 95% CI (-.085, -.006), further, acquisition (M= .174) didn’t differ significantly from extinction (M= .132) (p = .094, 95% CI (-.004, .089) however acquisition phase differed significantly from extinction recall (M= .116) (p= .019, 95% CI (.007, .110). Extinction (M= .132) didn’t differ significantly from extinction recall (M= .116) phase (p = 1.00, 95% CI (-.021, .053). To further explore the stimulus type x phase interaction, we performed a 2 x 2 repeated measures ANOVA with stimulus type (CS+, CS-) as within subject factor and group (IC-SE, SE) as between subject factor was conducted on SCR ratings for each phase separately. In habituation phase the results indicated non-significant main effect of stimulus type [F (1,49) = .078, p= .781, ƞp 2 =.002] indicating no difference in baseline physiological arousal in CS+ (M= .125, SD= .142) and CS- (M=.133, SD=.160) during habituation. Further, no significant stimulus x group interaction effect was observed. In acquisition phase the results indicated a significant main effect of stimulus type [F (1,49) = 17.27, p < .001, ƞp 2 =.261] with higher SCR for CS+ (M= .207, SD= .120) than CS- (M= .142, SD= .104) this indicates successfully fear acquisition. Further, no significant stimulus x group interaction effect was observed, implying comparative fear acquisition in both groups. In extinction phase the results indicated a non- significant main effect of stimulus type [F (1,49) = 2.26, p = .139, ƞp 2 = .044] this indicates physiological arousal to CS+ and CS- was comparable after extinction. Further, a non-significant stimulus x group interaction effect was observed. However, a pairwise comparison indicated a significant main effect of group (p = .006, 95% CI (.024, .132). An independent sample t-test was conducted to assess the group difference on CS+ and CS- SCR values, the results revealed a significant group difference on CS+ values t (49) = 2.21, p = .032, d= .621, with lower CS+ values in SE group (M =.107, SD=.118) than IC-SE group (M= .178, SD = .109). Further, it also revealed a significant group difference on CS- values t (49) = 2.998, p = .004, d= .841, with lower SCR values in SE group (M= .079, SD =.091) than IC-SE group (M= .164, SD = .109). This implies an overall increase in physiological arousal in IC-SE group. In extinction recall phase the results indicated a non-significant main effect of stimulus type [F (1,49) = 3.60, p =.064, ƞp 2 = .068]. Further, a non-significant stimulus x group interaction effect was also observed (Figure 5). Figure 5 SCR plots by (A) differential values and (B) CS Type Valence ratings We performed a 4 x 2 x 2 repeated measure ANOVA with phase (habituation, acquisition, extinction and extinction recall) and stimulus type (CS+. CS-) as within subject factor and group (IC-SE & SE) as between subject factor on valence ratings. The results indicated a significant main effect of phase [F (2.66, 130.35) = 3.98, p< .012, ƞp2 = .075; see Footnote 1], further, a significant main effect of stimulus type [F (1,49) = 29.29, p< .001, ƞp2 =.374]. Additionally, the results also indicated a significant phase x stimulus type interaction [F (2.44,119.47) = 8.89, p< .001, ƞp2 =.154; see Footnote 1]. Further, a pairwise comparison of phase revealed a significant difference between habituation (M= 5.25) and acquisition (M= 4.63) phase (p = .029, 95% CI (.05, 1.21), acquisition (M= 4.63) did not differ significantly from extinction (M= 5.18) (p= .159, 95% CI (-1.21,.11) but differed significantly from extinction recall (M= 5.39) (p = .031, 95% CI (-1.48, -.05). Further, extinction (M= 5.18) didn’t significantly differ from extinction recall (M= 5.39) phase (p = 1, 95% CI (-.76, .34). Additionally, a pairwise comparison of stimulus revealed a difference in CS+ (M= 4.42) and CS- (M= 5.80) (p< .001, 95% CI (-1.90, -.87)). To further explore the stimulus type x phase interaction, we performed a 2 x 2 repeated measures ANOVA with stimulus type (CS+, CS-) as within subject factor and group (IC-SE, SE) was conducted on valence ratings for each phase separately. In habituation phase the results indicated non-significant main effect of stimulus type [F (1,49) = .018, p= .895, ƞp 2 =.000] indicating no difference in baseline valence in CS+ (M= 5.235, SD= 1.976) and CS- (M=5.275, SD= 1.919) during habituation. Further, no significant stimulus x group interaction effect was observed. In acquisition phase the results indicated a significant main effect of stimulus type [F (1,49) = 28.006, p < .001, ƞp 2 =.364] with lower valence for CS+ (M= 3.490, SD= 2.148) than CS- (M= 5.784, SD= 2.194) this indicates successfully fear learning. Further, no significant stimulus x group interaction effect was observed, implying comparative fear acquisition in both groups. In extinction phase the results indicated a significant main effect of stimulus type [F (1,49) = 20.784, p < .001, ƞp 2 =.298] with lower valence for CS+ (M= 4.333, SD= 1.956) and CS- (M= 6.000, SD= 2.126). Further, a non-significant stimulus x group interaction effect was observed. In extinction recall phase the results indicated a significant main effect of stimulus type [F (1,49) = 20.594, p < .001, ƞp 2 =.296] with lower valence for CS+ (M= 4.647, SD= 1.874) and CS- (M= 6.118, SD= 2.094). Further, a non-significant stimulus x group interaction effect was observed (Figure 6). Figure 6 Subjective valence plots by (A) differential values (B) CS Type Fear ratings We performed a 4 x 2 x 2 repeated measure ANOVA with phase (habituation, acquisition, extinction and extinction recall) and stimulus type (CS+. CS-) as within subject factor and group (IC-SE & SE) as between subject factor on fear ratings. The results indicated a significant main effect of phase [F (2.46,121.28) = 12.85, p< .001, ƞp2 = .208; see Footnote 1] and a significant main effect of stimulus type [F (1,49) = 79.84, p< .001, ƞp2 =.620]. Additionally, the results also indicated a significant phase x stimulus type interaction [F (2.23, 109.29) = 31.06, p< .001, ƞp2 =.388; see Footnote 1]. Further, a pairwise comparison of phase revealed a significant difference between habituation (M= 3.04) and acquisition (M= 4.02) phase (p< .001, 95% CI (-1.52, -.44), acquisition (M= 4.02) differed significantly from extinction (M= 3.28) (p< .001, 95% CI (.29, 1.20) and extinction recall (M= 2.82) (p < .001, 95% CI (.65, 1.77). Further, extinction (M= 3.28) didn’t significantly differ from extinction recall (M= 2.82) phase (p = .064, 95% CI (-.02, .95). Additionally, a pairwise comparison of stimulus revealed a difference in CS+ (M= 4.34) and CS- (M= 2.23) (p< .001, 95% CI (1.63, 2.57)). To further explore the stimulus type x phase interaction, we performed a 2 x 2 repeated measures ANOVA with stimulus type (CS+, CS-) as within subject factor and group (IC-SE, SE) was conducted on fear ratings for each phase separately. In habituation phase the results indicated non-significant main effect of stimulus type [F (1,49) = .004, p=.950, ƞp 2 =.000] indicating no difference in baseline fear rating in CS+ (M=3.00, SD=1.929) and CS- (M= 3.019, SD=2.102) during habituation. Further, no significant stimulus x group interaction effect was observed. In acquisition phase the results indicated a significant main effect of stimulus type [F (1,49) = 93.313, p < .001, ƞp 2 =.656] with higher fear ratings for CS+ (M= 5.843, SD=2.411) than CS- (M= 2.177, SD=1.545) this indicates successfully fear learning. Further, no significant stimulus x group interaction effect was observed, implying comparative fear acquisition in both groups. In extinction phase the results indicated a significant main effect of stimulus type [F (1,49) = 51.481, p < .001, ƞp 2 = .512] with higher fear rating for CS+ (M= 4.549, SD=2.229) than CS- (M= 1.980, SD= 1.771). Further, a non-significant stimulus x group interaction effect was observed. In extinction recall phase the results indicated a significant main effect of stimulus type [F (1,49) =, 66.907 p < .001, ƞp 2 = .577] with higher fear ratings for CS+ (M= 3.682, SD=2.059) than CS- (M=1.725, SD= 1.358). Further, a non-significant stimulus x group interaction effect was observed (Figure 7). Figure 7 Subjective fear plots by (A) differential values (B) CS Type

The major findings of the current study are (i) inhibitory control training results in better reduction in UCS-expectancy during extinction and better recall of UCS- expectancy extinction during extinction recall, (ii) further, inhibitory control training results in lower subjective arousal ratings during extinction and sustained effect during extinction recall phase (iii) our results also provide support to retrieval stopping hypothesis of extinction. Our findings show that training on stop-signal task led to better reduction in UCS- expectancy in experimental group (IC-SE), this implies a positive relation between inhibitory control and inhibitory learning. This suggests that a general increase in inhibitory control enhances inhibitory learning contributing to better formation of inhibitory association between stimuli during extinction. Our findings support the notion that inhibitory learning is a central process of extinction (Bouton, 1993; Wagner, 1981). To the best of our knowledge the current study is the first investigation that focusses on enhancing extinction through training individual capacity to form inhibitory associations, our results imply inhibitory control plays a role in process of extinction. Further, we also found enhanced reduction in subjective arousal in IC-SE group. This could be explained as a result of reduced UCS-expectancy during extinction. Since the UCS-expectancy is low during extinction i.e. the participants’ anticipation to encounter the UCS is low, this could have resulted in subjective arousal associated with the CS+ to be reduced as well. Although, we observed significant difference between groups in UCS-expectancy and subjective arousal, we failed to observe the same result in subjective valence and fear ratings. This could be explained through evaluative conditioning, the CS+ valence is not entirely acquired through CS-UCS association (Hofmann et al., 2010). Inhibitory control training resulted in reduced UCS-expectancy through better inhibitory learning i.e. CS- no UCS association, however it did not change the negative evaluation associated with UCS (Davey, 1989). Therefore, we did not find any difference on valence and fear ratings between the groups. We observed an increased SCR to both CS+ and CS- in IC-SE group in comparison to the SE. Stop-signal task requires constant monitoring and attention to the go and the stop-signals and also responding to the go signals within a time limit. Previous studies have shown increase in physiological arousal during inhibitory control tasks. (Hajcak et al., 2003). Stop-signal task was immediately followed by extinction which could have resulted in a heightened arousal in the IC-SE group. Although, SCR values in the experimental group reduced from acquisition to extinction showing effect of extinction learning, however in comparison to the control group participants exhibited a higher SCR value for both CS+ and CS- which could be attributed to the performance of inhibitory control task. The increased SCR in IC-SE group could also be explained in terms of emotional interference in inhibitory control. Participants trained on inhibitory control task resulted in better formation of inhibitory associations during extinction as evidenced through significantly reduced UCS expectancy in comparison to the control group. However, it might have occurred that the newly enhanced inhibitory control might have resulted in more cognitive effort in forming the inhibitory association due the emotional stimuli involved in extinction, hence resulting in increased physiological arousal. Previous research has shown that emotional stimuli interfere with inhibitory control (Verbruggen & De Houwer, 2017) that could have resulted in increased cognitive effort, further resulting in changes in skin conductance response (Westbrook & Braver, 2015). Future research should explore the effect of emotional stop-signal task on extinction learning, perhaps that might result in lower cognitive effort in updating inhibitory associations and hence result in reduced SCR. In our results we also observed better extinction recall of UCS-expectancy and arousal ratings in experimental group. This result can be explained by better consolidation of extinction memory on day 2. Previous studies have demonstrated that under heightened arousal pre- and post- learning results in better memory consolidation. Previous work has shown that cortisol administration before presentation of negative and neutral images resulted in better recall of emotional images in comparison to a placebo group (Buchanan and Lovallo, 2001; Kuhlmann & Wolf, 2006). Bentz and colleagues, 2013 demonstrated that stress prior to extinction results in better retrieval of UCS-expectancy on subsequent day. We found similar results of reduced UCS- expectancy ratings during extinction recall. However, in our study we also found better reduction in UCS-expectancy during extinction as well, this could be due to the inhibitory control task which might have resulted in increased arousal but also resulted in overall increase in inhibitory learning. Previous studies have studied effect of heightened arousal on memory through epinephrine or cortisol administration (Cahill & Alkire, 2003; Buchanan and Lovallo, 2001; Kuhlmann & Wolf, 2006), negative physiological stress (cold pressor) (Cahill et al., 2003), negatively arousing stimuli (Cahill & McGaugh, 1995), negatively stressful and positively arousing videos (Liu, Graham & Zorawski, 2008). These studies aimed to study the effect of heightened arousal on memory consolidation and explicitly manipulated arousal and used appropriate measurement of arousal resulting from the manipulation. In our study we didn’t explicitly intend to manipulate arousal, therefore no measures for induction of arousal were used. However, given the nature of the task performed by the IC-SE group, it could have resulted in heightened arousal, and further maintained due the presentation of arousing stimuli in extinction phase consequently resulting in better memory consolidation. Our results suggest induction of heightened arousal before extinction results in better consolidation of extinction learning, this is in line with the previous study suggesting that arousal induced before learning can enhance the memory consolidation even after 30 minutes of arousal induction (Tambini et al., 2017). Further, our results provide support to retrieval stopping hypothesis of extinction. Both retrieval stopping and action stopping involve activation of right dorsolateral and ventrolateral prefrontal cortexes, training on stop-signal task could have resulted in domain-general improvement in inhibition resulting in better extinction. This implies potential involvement of retrieval stopping in extinction. Additionally, our results also suggest a domain general nature of inhibitory control involved in both inhibition of motoric and inhibition involved in extinction (nonmotor processes). Further, Anderson & Floresco (2022) suggest that inhibitory control during extinction aids in formation of inhibitory association which after consolidation works in harmony with or substitutes inhibitory control, resulting in better extinction and further better retention of extinction in IC-SE group. In IC-SE group due to stop-signal task training, inhibitory control might play a more active role in aiding extinction and hence resulting in better extinction recall.

Our results highlight that differences on inhibitory control might be one factor that could lead to variations on extinction learning. Further, training on inhibitory control task results in better inhibitory learning. This can be of application to individuals with anxiety disorders, who show deficits in inhibitory learning. Training such individuals on inhibitory control task might aid exposure therapy outcome. Further, improvement in extinction through stop-signal task implies involvement of similar inhibitory mechanisms in both processes.

and future direction In our study the inhibitory control training was of shorter duration, future research could investigate the effect of longer duration of training inhibitory control on extinction and spontaneous recovery. Longer training duration might result in reduced physiological responses as well. Further, including another group trained on emotion stop-signal task might have result in decreased SCR during extinction, this can be explored in future studies. Additionally, we trained individuals on stop-signal i.e. response inhibition, whether other training on inhibitory control task like Stroop task or flanker task result in similar effect on extinction needs to be investigated. Further, the control group did not perform any task therefore it remains uncertain whether any other task might have resulted in similar higher arousal consequently resulting in better extinction recall. Funding statement The corresponding author is supported by the F.I.G. grant (IITR/SRIC/2741). The funding agency had no role in the preparation of the manuscript. Ethical approval The project was approved by the Institute Human Research Ethics Committee, Indian Institute of Technology Roorkee (IITR/ICC/24/18). Availability of Data The data will be made available upon request from the corresponding author, the study was not preregistered. Code availability The code will be made available on request from the corresponding author. Acknowledgments We would like to thank the participants for filling out the questionnaires. The authors also thank the Memory and Anxiety Research Group (MARG), Indian Institute of Technology Roorkee for its constant support. Clinical trial registration Not applicable Author Contribution Kaneez Fatima Dar : Conceptualization; Data curation; Formal analysis; Investigation; Methodology; Project administration; Software; Visualization; Writing—original draft; Writing— review and editing Manish Kumar Asthana : Conceptualization; Formal analysis; Investigation; Methodology; Funding acquisition; Resources; Validation Writing— review and editing.

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Clinical Psychological Science, 10 (4), 622–639. https://doi.org/10.1177/21677026211055169 Information & Authors Information Version history Peer review timeline Published International Journal of Psychophysiology Version of Record1 Mar 2026Published Copyright This work is licensed under a Non Exclusive No Reuse License. Authors Metrics & Citations Metrics Article Usage 367views 173downloads Citations Download citation Kaneez Fatima Dar, Manish Kumar Asthana. Inhibitory control training might be a gateway to enhance fear extinction.. Authorea. 24 January 2025. DOI: https://doi.org/10.22541/au.173773024.42679517/v1 DOI: https://doi.org/10.22541/au.173773024.42679517/v1 If you have the appropriate software installed, you can download article citation data to the citation manager of your choice. Simply select your manager software from the list below and click Download. For more information or tips please see 'Downloading to a citation manager' in the Help menu.