The time course of affective processing in anhedonia: insights from event-related potentials

The time course of affective processing in anhedonia: insights from event-related potentials | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Article The time course of affective processing in anhedonia: insights from event-related potentials Valentina Mologni, Carola Dell’Acqua, Roza Mejza, Simone Messerotti Benvenuti This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7073207/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Anhedonia, defined as a diminished capacity to experience pleasure form appetitive stimuli, is a core symptom of depression and a predisposing factor for its development. Prior research links anhedonia with blunted emotional processing of appetitive stimuli. Yet, emotional processing encompasses multiple stages (cue engagement, affective anticipation, and elaboration), and how each stage relates to anhedonia remains unclear. This study examined these associations in a sample of university students ( n = 45, 31 females) with varying levels of anhedonia. Participants underwent electroencephalography recording during an S1-S2 paradigm, in which a cue (S1) anticipated the valence (pleasant, neutral, unpleasant) of an upcoming emotional image (S2). Three event-related potentials (ERPs) were assessed: the Cue-P300 (reflecting cue evaluation and engagement), the Stimulus Preceding Negativity (SPN; reflecting affective anticipation), and the Late Positive Potential (LPP; reflecting affective elaboration). While the LPP was larger for emotional vs. neutral images, the Cue-P300 and the SPN were more pronounced for pleasant (but not unpleasant) vs. neutral stimuli. Notably, anhedonia, independent of other depressive symptoms, was associated with an increased SPN and a blunted LPP for pleasant stimuli. These findings suggest a complex pattern of emotional processing in anhedonia, marked by increased anticipation but reduced elaboration of appetitive stimuli. Health sciences/Diseases Biological sciences/Neuroscience Biological sciences/Psychology Social science/Psychology Anhedonia depression vulnerability ERPs S1-S2 task SPN LPP Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction Depression is recognized as a leading cause of disability, affecting over 300 million people globally [ 1 ]. It manifests through a range of affective, cognitive, and somatic symptoms [ 2 , 3 ]. Given its substantial burden, identifying viable factors contributing to depression vulnerability has been highlighted as a core priority [ 4 ]. In this regard, emerging research suggests that anhedonia—defined as the loss of pleasure and emotional disengagement from appetitive stimuli—may play a central role in the disorder's onset and maintenance, being a potential endophenotype for depression [ 5 ]. Notably, anhedonia can precede the onset of depression and often persists even after other symptoms have remitted [ 6 ]. However, anhedonia is not a unitary construct, and addressing its heterogeneity could enhance our understanding of depression vulnerability [ 7 ]. Aligned with this purpose, the Research Domain Criteria (RDoC) initiative proposed a dimensional model whereby integrating multiple levels of analysis of different constructs might be useful to fully characterize anhedonia. Among the RDoC constructs, the Positive Valence Systems (PVS) may be particularly relevant for its understanding, as they encompass a set of systems involved in anticipating, obtaining, elaborating, and responding to pleasant or rewarding stimuli [ 8 , 9 ]. While these phases are related, they are also dissociable at both behavioral and neural levels, each of them serving distinct functions: anticipation allows individuals to prepare for future events, while elaboration and response mobilization entail engaging with and processing the immediate reward. Reduced appetitive anticipation and elaboration has been observed in depression [ 10 , 11 , 12 , 13 ], in children with familial risk of depression [ 14 ], and it can prospectively predict its onset [ 15 , 16 , 17 ]. Yet, it is still unclear whether depression vulnerability stems from deficits occurring at any specific stage of appetitive stimulus processing [ 18 , 19 ]. For example, anhedonia may not necessarily reflect a diminished ability to experience pleasure, but rather a deficit in mobilizing motivational resources to pursue pleasurable activities [ 20 ]. Studies employing functional magnetic resonance imaging (fMRI) helped disentangle the existence of both shared and dissociable neural mechanisms related to distinct phases of appetitive stimulus processing [ 21 , 22 ]. Specifically, the anticipation of appetitive stimuli consistently engages the ventral striatum, while the elaboration phase is related with heightened activity in the orbitofrontal cortex [ 23 , 24 , 25 ] and regulatory involvement of dorsolateral and ventromedial prefrontal cortices [ 26 ]. Notably, altered cortical and striatal activation during the anticipation and processing of appetitive stimuli has been observed in adults with current depression [ 27 , 28 , 29 ], in remission [ 30 ], and in youths at risk for depression [ 31 ]. However, most of these studies have relied upon monetary reward tasks, without examining the distinct phases of other appetitive stimuli processing (e.g., pleasant images, social rewards) that might be more relevant to the anhedonic symptomatology in the context of depression risk [ 32 , 33 ]. For example, greater neural responses to pleasant pictures but not monetary rewards have been associated with improved treatment outcomes for depression [ 34 ]. More importantly, due to its poor temporal resolution, fMRI might be inadequate for the online tracking of the distinct phases of appetitive stimulus processing, leading to the merging of neural activity associated with processes that are temporally close but psychologically distinct (i.e., anticipation, stimulus response, elaboration) [ 35 ]. These issues can be addressed by using event-related potentials (ERPs) of the electroencephalographic (EEG) signal [ 36 ]. Thanks to their excellent temporal resolution, ERPs allow for the precise examination of distinct phases of appetitive stimulus processing. An extensively used paradigm in ERP research to tackle the anticipation and elaboration of affective stimuli is the S1-S2 task, which involves the presentation of a series of paired trials in which an initial cue stimulus (S1) indicates the emotional valence of a second stimulus (S2, or imperative stimulus) that appears after a fixed inter-stimulus interval (ISI) [ 37 , 38 , 39 ]. This allows the dissociation of anticipation from the elaboration of appetitive stimuli in the brain. The Cue-P300 and the Stimulus Preceding Negativity (SPN) are two main components associated with anticipation, whereas elaboration is associated with the Late Positive Potential (LPP). The Cue-P300 is a positive parietal deflection occurring at about 300 ms after cue (S1) onset, which reflects affective engagement (i.e., the allocation of attentional resources engaged in the upcoming stimulus) towards the upcoming stimulus (S2) and is typically larger for emotional vs. neutral stimuli [ 40 ]. In addition, the Stimulus Preceding Negativity (SPN) is a slow negative potential that emerges approximately 200 ms before the S2, exhibiting a larger amplitude (i.e., a greater negativity) during the anticipation of emotional vs. neutral stimuli at frontocentral sites [ 41 , 42 ]. The SPN is linked to the anticipation of emotionally arousing content [ 43 ] and is modulated by the level of motivational engagement with the forthcoming stimulus. Finally, the LPP is a positive waveform that begins approximately 400 ms following the S2 onset, with maximal amplitude at centro-parietal sites. This ERP reflects elaborative processing, including motivated attention, as well as stimulus representation and evaluation [ 44 ]. In addition, emotional stimuli consistently elicit a larger LPP compared to neutral stimuli [45] reflecting the activation of the appetitive motivational system in the brain [ 46 ]. Investigating whether individuals who are vulnerable to depression – such as those experiencing anhedonic symptoms – exhibit affective processing alterations which are analogous to those observed in individuals with clinical depression may help identify early indicators of the disorder and clarify whether these alterations are potential precursors rather than mere characteristics of the clinical condition. Consistently, research on affective processing has reported a blunted LPP to pleasant stimuli in not only in individuals with diagnosed depression [ 47 , 48 ], but also in those at risk for the disorder, such as individuals with subclinical symptoms or familial risk for depression [49,50,51]. This finding suggests that reduced affective elaboration of pleasant content, indexed by a blunted LPP amplitude, may serve as a psychophysiological marker of depression vulnerability, potentially reflecting anhedonia [ 48 ]. In line with this, the limited number of studies examining the two ERP components related to anticipation (i.e., Cue-P300 and SPN) have found a reduction in the Cue-P300 amplitude in response to rewarding stimuli among individuals with both subclinical and clinical depression [ 52 , 53 ]. Interestingly, blunted Cue-P300 has been observed in individuals with anhedonia but without current depressive symptoms during a gambling task [ 54 ], suggesting that anhedonia may influence affective engagement for future rewards. In contrast, studies using reward tasks have yielded mixed findings regarding the SPN, likely due to differences in task design and that this component is sensitive to several manipulations [38, 55, 56, 57]. For instance, the SPN amplitude has been reported as blunted [ 58 ], increased [59], or unchanged [ 52 ] in individuals with depressive symptoms relative to controls. Moreover, two studies using reward tasks and assessing anhedonia while controlling for depressive symptoms reported no significant change in SPN amplitude during the anticipation of rewarding relative to non-rewarding stimuli [ 54 , 60 ]. Given that most of the ERPs research has relied on traditional reward-based paradigms — primarily assessing the processing of external incentives such as monetary rewards — the S1–S2 task enables the investigation of intrinsic affective responses to emotionally salient stimuli, such as pleasant images [ 37 , 38 ]. This distinction is particularly important, as heightened neural responses to pleasant images – but not to monetary rewards – have been associated with more favorable treatment outcomes in individuals with depression [ 34 ]. In addition, the temporal dynamics of affective processing of pleasant stimuli remain underexplored, as most existing studies have focused exclusively on the elaboration stage [ 47 , 48 , 49 , 50 ]. The reviewed literature suggests that appetitive stimuli processing might contribute to anhedonia and, consequently, depression vulnerability. However, there is still limited evidence examining the link between anhedonia and the distinct appetitive picture processing stages. Hence, this study aimed to examine appetitive picture processing through the assessment of ERPs during an emotional S1-S2 task in a sample of young adults with varying levels of anhedonia. Based on the reviewed literature, it was hypothesized that higher levels of anhedonia would be associated with reduced affective elaboration of pleasant stimuli, as indicated by a blunted LPP amplitude. Moreover, although the literature is still scarce, it was hypothesized that higher levels of anhedonia would be associated with reduced appetitive cue engagement and anticipation, as indexed by decreased amplitude of Cue-P300 and SPN components. While the primary focus was on appetitive processing in anhedonia, an exploratory analysis of the affective processing of unpleasant stimuli was also conducted. Lastly, this study explored whether the observed appetitive processing dynamics are uniquely associated with anhedonia or extend to non-anhedonic depressive symptoms. Methods Participants Forty-five Italian Caucasian Students (31 females, mean (M) age = 21.7 years, standard deviation (SD) = 1.84 years, range = 19–26 years) from the University of Padua (Italy) voluntarily participated in the study. An a-priori power analysis was performed using G*Power 3 [ 61 ]. According to methodological guidelines suggesting that ERPs are adequately powered to detect large effects [ 62 ], the power analysis indicated that a sample size of 41 participants would be sufficient to detect a medium-to-large effect size (f = 0.3) with a statistical power of 0.8 (α = 0.05, predictors = 3). The enrolled sample was medically healthy and free from psychotropic medication, as assessed with an ad-hoc interview. Exclusion criteria included a current diagnosis or history of cardiovascular and neurological diseases and a formal diagnosis of mental diseases. All participants had a normal-to-corrected vision and were naïve to the purpose of the experiment. All participants understood and signed the informed consent forms. The study was conducted in compliance with the Declaration of Helsinki on research on human subjects and was approved by the Ethical Committee of Psychological Research, Area 17, University of Padua (prot. no. 564-b). Participants received a monetary compensation of 25 euros for their participation. Psychological measures The Snaith-Hamilton Pleasure Scale (SHAPS) was used to assess anhedonia (internal consistency: Kuder-Richardson coefficient = 0.85) [ 63 ]. The SHAPS consists of 14 items that cover four domains of pleasure experience: interest, social interaction, sensory experience, and food/drink. Each item (e.g. I would be able to enjoy a beautiful landscape or view) is rated on a 4-point scale from “strongly disagree” to “strongly agree.” Responses of “disagree” score 1 point each, while “agree” responses score 0 point each. The total score ranges from 0 to 14, with scores of 3 or higher indicating a reduced ability to experience pleasure, and consequently, the presence of anhedonic symptoms. To evaluate depressive symptoms over the past two weeks, the Italian version of the Beck Depression Inventory-II [ 64 , 65 ] was administered. This self-report questionnaire comprises 21 items, each rated on a four-point Likert scale, with total scores ranging from 0 to 63 (internal consistency: Cronbach's alpha = 0.87). Higher scores indicate more severe depressive symptoms. In the Italian version, a score of 12 is the cut-off point for distinguishing between individuals with and without depressive symptoms [ 65 ]. From the BDI-II, a non-anhedonic symptom (sum of all items except 4, 12, 22) subscale was derived (internal consistency: Cronbach's alpha = 0.83) [ 66 ]. Experimental task In this study, participants engaged in an S1-S2 emotional paradigm while undergoing electroencephalographic (EEG) recording. The task consisted of a single block of 84 trials. Each trial started with a 500 ms baseline (a white fixation dot), followed by a cue (S1) lasting 1500 ms that signaled the emotional content (a plus for pleasant, a circle for neutral, a minus for unpleasant) of the upcoming picture (S2), presented after the 4500 ms inter-stimulus interval (ISI) and lasting 6000 ms. The S2 was followed by a variable inter-trial interval (ITI) of 6000–8000 ms, during which a white fixation dot (identical to the baseline) was presented (see Supplementary, Figure S1 online). Participants were instructed to observe the cue (S1) and the subsequent emotional image (S2), with no motor response required. They were also informed that the emotional image would always be preceded by a congruent cue, which would anticipate its emotional valence. The S2 stimuli comprised 42 emotional images (600 x 800 pixels), each repeated twice, selected from the International Affective Picture System [ 67 ] including 14 pleasant images (i.e., erotic couples, sports), 14 neutral images (i.e., neutral scenes, household objects), and 14 unpleasant images (i.e., attacking humans, explosions, animals). The selection was based on normative arousal ratings, ensuring that pleasant and unpleasant images were matched in arousal levels (pleasant, mean ± SD = 6.5 ± 0.3; unpleasant, mean ± SD = 6.2 ± 0.6; p = .11) which were significantly higher than those of neutral pictures (neutral, mean ± SD = 3.1 ± 0.5, all ps < .001) [ 68 ]. To induce greater psychophysiological changes, only highly arousing pleasant and unpleasant pictures were selected [ 69 ]. Stimuli were presented in a semi-randomized sequence to prevent consecutive S1-S2 pairs from having the same emotional valence. At the end of the task, the 42 images (14 for each emotional category) shown during the S1-S2 paradigm were presented again, and ratings of emotional valence and arousal were obtained using a computerized version of the 9-point Valence and Arousal scales of the Self-Assessment Manikin (SAM) [70]. (IAPS catalogue numbers: 1120, 1201, 1301, 1525, 2493, 2495, 2512, 2575, 2595, 2840, 4611, 4647, 4651, 4652, 4670, 4695, 5532, 6200, 6250, 6260, 6312, 7002, 7004, 7009, 7036, 7041, 7224, 7547, 8034, 8080, 8180, 8185, 8200, 8370, 8400, 8490, 9185, 9622, 9630, 9635.2, 9909, 9940). Procedure The study was advertised via various means, including flyers and online platforms. Interested students from the University of Padua completed an online form, administered through Google Forms. The form encompassed an anamnestic interview to assess eligibility, the SHAPS, and the BDI-II. Eligible participants were invited to a laboratory session in the Clinical Psychophysiology laboratory at the Department of General Psychology of the University of Padua. Participants were instructed to refrain from alcohol consumption on the day before the experimental session and to avoid caffeine and nicotine for at least four hours before the session. Upon arrival at the laboratory, participants read and signed the informed consent form, after which they underwent an ad-hoc anamnestic interview. Participants were then seated on a comfortable chair in a dimly lit, sound-attenuated room. This study was part of a broader project that also included the recording of the electrocardiogram (ECG), in addition to the EEG. After electrode attachment and a 3-minute resting-state recording, they were given three practice trials, one for each emotional category (pleasant, neutral, unpleasant). Following the practice trials, participants completed the S1-S2 paradigm. The entire procedure took approximately 90 min. Electroencephalogram data acquisition and analysis EEG was recorded using a 32-channel ANT system and a computer running eego™ software (ANT Neuro, Enschede, Netherlands). The elastic cap with 32 tin electrodes was arranged according to the 10–20 System (FP1, FPz, FP2, F7, F3, Fz, F4, F8, FC5, FC1, FC2, FC6, T7, C3, Cz, C4, T8, CP5, CP1, CP2, CP6, P7, P3, Pz, P4, P8, POz, O1, Oz, O2, and M1 and M2 [mastoids]), referenced online to CPz. Vertical and horizontal electrooculograms (EOGs) were recorded using a bipolar montage to track horizontal eye movements and blinks, with electrodes placed above and below the right orbit and at the external canthi of the eyes. Electrode impedance was kept below 10 kΩ. The EEG and the EOG signals were recorded in DC with a low-pass filter of 30 Hz and sampled at 1000 Hz. EEG data was resampled to 500 Hz and re-referenced offline to a linked mastoids montage in EEGLAB [ 71 ]. Further analyses were performed in Brainstorm [ 72 ]. Data was band-pass filtered from 0.01 to 30 Hz and corrected for blink artifacts using Independent Component Analysis (ICA). The EEG signal was segmented into 8500 ms epochs, from 500 ms before S1 to 2000 ms after the onset of S2 (i.e., -500 to 8000 ms). Based on previous research, the signal was baseline-corrected using the average signal activity within the − 250 ms to -50 ms time window preceding S1. This window was selected for each ERP, as it was the period of EEG recording being closest to the participant's true baseline, without other cognitive or emotional processes occurring [ 38 , 73 ]. A semiautomatic procedure was employed for artifact rejection, applying a voltage difference threshold of 200 µV (peak-to-peak). Then, epochs were visually inspected to detect and reject possible remaining artifacts. According to previous research [38,74,75] and the visual inspection of the grand-average ERPs waveforms for the three emotional categories, the Cue-P300 and the LPP were maximal at parietal sites (Fig. 1 ), while the SPN was prominent at frontocentral sites (Fig. 2 ). Therefore, the Cue-P300 was calculated by averaging the peak amplitudes at the parietal poll (P3, PZ, P4) in the time window from 200 to 400 ms post-S1, while the SPN was calculated as the mean amplitude at the frontocentral poll (FC1, FC2, FC5, FC6) within the 200 ms preceding the onset of S2. Further, the LPP was calculated by averaging the mean amplitude at the parietal poll (P3, PZ, P4) in the time window from 500 to 1000 ms post-S2. The grand average ERP waveforms for parietal, central, and frontal electrodes during the S1-S2 task, are presented in the supplementary material (see Supplementary, Figure S2 online). Statistical Analyses Statistical analyses were conducted in Rstudio [ 76 ]. A p -value of .05 was used as the threshold for statistical significance. Experimental hypotheses were tested using linear mixed-effect models with the lme4 and lmerTest packages [ 77 , 78 ]. All models included participants as a random intercept, and sex (biological) as a covariate to account for the sex unbalance within the experimental sample. To test the effect of emotional category and anhedonia on self-reported ratings of valence and arousal, two separate models with Category (i.e., pleasant, neutral, unpleasant), SHAPS scores, and their interaction as fixed factors were conducted: Model ← lmer(Subjective rating of arousal or valence ∼ Category × SHAPS + Sex + (1|Subject)). Then, to ensure the effect of emotional valence on the Cue-P300, SPN, and LPP, three separate models with Category as a fixed factor were conducted on each ERP: Model ← lmer(ERP amplitude ∼ Category + Sex + (1|Subject)). To test the primary hypothesis—examining the relationship between anhedonia levels and distinct stages of emotional processing—three separate models were conducted, one for each ERP component (Cue-P300, SPN, LPP). Differential ERP scores (ΔERP; pleasant - neutral, unpleasant - neutral) were employed to reduce the number of predictors and isolate the effect of the emotional category [ 79 ]. The models included Category (pleasant, unpleasant), SHAPS scores, and their interaction as fixed factors: Model ← lmer(△ERPs amplitude ∼ Category × SHAPS + Sex + (1|Subject)). To test whether the examined effects are uniquely related to anhedonic symptoms, the same models were conducted with non-anhedonic depressive symptoms as predictor. Particularly, Category (pleasant, unpleasant), BDI-II non-anhedonic scores, and their interaction were included as fixed effects in the model: Model ← lmer(△ERPs amplitude ∼ Category × BDI-II non-anhedonic + Sex + (1|Subject)). For the fixed effects, the estimated coefficients ( b ), standard errors (SE), t values, and confidence intervals for each parameter were reported. Additionally, p -values obtained through the Satterthwaite approximation, as implemented in the lmerTest package [ 78 ], were provided. All predictors were centered and scaled. The collinearity was tested by calculating the Variance Inflation Factors (VIF) with the vif function of the car package [ 80 ]. Significant categorical effects ( p < .05) were followed by Tukey HSD post-hoc tests to correct for multiple comparisons. Results Psychological measures and valence and arousal self-report ratings The average SHAPS score was 1.8 (SD = 1.7, range = 0–6), with 16 participants (35.55% of the total sample) scoring above the cut-off (i.e., ≥ 3). The average BDI-II score was 13.82 (SD = 10.11, range = 1–45), with 24 participants (53.33 % of the total sample) scoring above the cut off (i.e., ≥ 12). Additionally, the average BDI-II non-anhedonic score was 12.22 (SD = 8.86, range = − 39). Linear mixed-effects models on self-reported SAM ratings revealed a significant main effect of Category in predicting valence (F (2,1843) = 907.15, p < .001) and arousal (F (2,1843) = 517.12, p < .001). Specifically, subjective ratings of valence were significantly higher for pleasant images compared to neutral and unpleasant images (all ps Tukey < .001), and significantly lower for unpleasant images compared to neutral images ( p Tukey < .001). In addition, subjective ratings of arousal were significantly higher for both pleasant and unpleasant compared to neutral images (all ps Tukey .09) emerged. Table 1 shows the means and standard deviations of SAM valence and arousal ratings. Table 1 Mean and standard deviation of valence and arousal ratings for pleasant, neutral, and unpleasant pictures in the current sample. SAM ratings Pleasant Neutral Unpleasant Valence 6.78 ± 1.18 5.20 ± 1.30 3.17 ± 1.65 Arousal 4.94 ± 2.37 2.47 ± 2.36 5.26 ± 1.68 Note . Data are M ± SD; SAM = Self-Assessment Manikin. The influence of emotional category on ERPs amplitude The mixed-effect models predicting the ERPs amplitude from the emotional Category revealed a significant effect of Category across all the ERPs (Cue-P300, F (2,358) = 10.27, p < .01; SPN, F (2,487) = 3.18, p = .04; LPP, F (2,358) = 21.17, p < .01). Table 2 shows the mean ERPs amplitude for each emotional Category. In the Cue-P300 model, the peak amplitude was larger for pleasant relative to neutral and unpleasant trials (all ps Tukey < .01), while no significant difference emerged between unpleasant and neutral trials ( p Tukey = .21). In the SPN model, the mean amplitude was larger (i.e., more negative) for pleasant relative to neutral trials ( p Tukey = .03), while no significant difference emerged between unpleasant relative to pleasant or unpleasant relative to neutral trials (all ps Tukey > .24). Finally, the LPP mean amplitude was larger for emotional (i.e., pleasant, unpleasant) relative to neutral trials (all ps Tukey < .01), while no significant difference between pleasant and unpleasant trials emerged ( p Tukey = .18). Figure 1 shows the grand averages of the Cue-P300 and LPP, while Fig. 2 shows the grand average of the SPN for the three emotional categories. Table 2 Mean and standard deviation of the ERPs amplitude (µV) across emotional Categories. ERPs amplitude (µV) Cue-P300 SPN LPP Category Pleasant 9.74 ± 4.06 1.79 ± 5.15 9.92 ± 7.09 Neutral 8.59 ± 3.50 3.08 ± 6.62 7.30 ± 6.79 Unpleasant 9.02 ± 3.84 2.62 ± 5.95 10.94 ± 8.97 Note . Data are M (SD). Anhedonia and the distinct phases of appetitive picture processing Table 3 shows the results of the mixed-effect model predicting ERPs amplitude from Category, SHAPS, and their interaction. VIF values were all < 1.22, indicating acceptable levels of multicollinearity among the predictor variables. Table 3 Linear mixed-models predicting △ERPs amplitudes from emotional category (pleasant, unpleasant) and SHAPS scores. Predictor b (SE) t p Cue-P300 model Category 0.72 (0.25) 2.87 < .01*** SHAPS 0.60 (0.39) 1.51 .13 Sex 0.70 (0.81) 0.86 .39 Category × SHAPS 0.17 (0.25) 0.68 .49 SPN model Category -0.84 (0.57) -1.47 .14 SHAPS 0.13 (0.91) 0.14 .89 Sex 0.19 (1.88) 0.10 .91 Category × SHAPS -1.62 (0.58) -2.80 < .01*** LPP model Category -1.02 (0.64) -1.60 .11 SHAPS 0.56 (0.81) 0.69 .49 Sex -1.26 (1.60) -0.78 .43 Category × SHAPS -1.99 (0.64) -3.11 < .01*** Notes. The reference levels are Category-unpleasant and Sex-females. Significant results are shown in bold, with asterisks indicating the level of statistical significance (* p < .05, ** p < .01, *** p < .001). The model predicting Cue-P300 peak amplitude showed an effect of Category (i.e., the mean amplitude was larger for pleasant vs. unpleasant images), but no other significant effect emerged. The model predicting the SPN amplitude revealed a significant interaction between Category and SHAPS, showing that higher levels of anhedonia were associated with increased (i.e., more negative) SPN to pleasant stimuli compared to unpleasant ones (Fig. 3 ). In addition, the model predicting LPP amplitude revealed a significant interaction between Category and SHAPS, indicating that higher levels of anhedonia were associated with reduced LPP to pleasant stimuli compared to unpleasant ones (Fig. 4 ). The results from the models predicting ERPs amplitude from Category, non-anhedonic BDI-II scores and their interaction are reported in the Supplementary Material (see Supplementary, Table S1 online). No significant effect of non-anhedonic BDI-II or Category × non-anhedonic BDI-II emerged. Discussion The present study examined whether anhedonia influenced the emotional processing of pleasant stimuli at different stages, as assessed with different ERPs during the emotional S1-S2 task. While anhedonia is traditionally described as the inability to feel pleasure, recent perspectives suggest it may be also linked to impairments in motivation and anticipation, rather than solely a lack of pleasure [ 7 , 81 ]. In this context and given that emotional processing involves multiple stages (i.e., anticipation, elaboration), understanding how each stage relates to anhedonia may be relevant for the early identification of specific processes involved in depression vulnerability. Notably, in the current sample, higher levels of anhedonia, but not non-anhedonic depressive symptoms, were associated with larger anticipation (SPN) and blunted elaboration (LPP) of pleasant stimuli. The results from the LPP are in line with the formulated hypothesis and suggest a reduced elaboration and motivated attention to pleasant stimuli in individuals with higher levels of anhedonia. This finding is consistent with prior research, which has reported blunted LPP in individuals with both clinical [ 48 ] and subclinical depression [ 68 ], among those with a familial risk for the disorder [ 50 , 82 ], and as a predictor of future depression onset [83]. Notably, in this study, a reduced LPP to pleasant stimuli was specifically associated with anhedonia rather than non-anhedonic depressive symptoms, suggesting that this component may uniquely reflect anhedonia rather than general depressive symptomatology, at least in individuals without a formal diagnosis of depression. Contrary to the initial hypothesis, the results from the SPN suggest that anhedonia may increase the anticipation of pleasant stimuli. This heightened anticipation could serve as a compensatory mechanism to counterbalance the already diminished elaboration and processing of these stimuli. Instead, in clinical depression both anticipation and elaboration may become impaired, reflecting a more extended deficit of appetitive stimuli processing [ 27 , 28 ]. Notably, a similar pattern of heightened anticipation has been observed in individuals remitted from a major depressive disorder [ 84 ] and in those exhibiting depressive symptoms [59] during reward processing tasks. Thus far, this is the only study highlighting this effect during an emotional picture processing paradigm, and further research employing similar tasks may aid in clarifying this pattern as a marker of depression vulnerability as well as its relationship with anhedonia. However, caution is warranted in interpreting this result, as the SPN, typically identified as a slow negative waveform, consistently maintained its amplitude above the isoelectric line across all emotional categories, possibly because the employed paradigm did not activate the appropriate anticipation mechanisms. Furthermore, the valence of the emotional stimuli was always predicted by corresponding cues, and this task might not have sufficiently engaged anticipatory mechanisms during the processing of emotional stimuli. The choice of maintaining congruency between S1 and S2 was made to ensure coherence with previous studies [ 37 , 38 , 43 ] but future research should consider incorporating such manipulations to better understand the dynamics of anticipation and emotional processing. In addition, neither anhedonia nor non-anhedonic depressive symptoms significantly predicted changes in Cue-P300 amplitude, indicating that these conditions did not influence the initial affective engagement and preparation to elaborate the emotional stimuli. This finding is in contrast with other studies reporting a reduced Cue-P300 in individuals with anhedonia [ 54 ] or depression vulnerability [ 52 ]. The discrepancy may stem from differences in experimental paradigms, as prior studies assessing the Cue-P300 often employed reward tasks that typically required a motor response. Taken together, the larger SPN and smaller LPP to pleasant images suggest a complex pattern of affective processing in individuals with anhedonia, marked by a rapid transition from increased anticipation to reduced elaboration specifically of pleasant stimuli. Although some studies have reported diminished affective responses to both unpleasant and pleasant stimuli in individuals vulnerable to depression [ 50 , 82 ], indicating a general reduction in motivational drive [ 85 ], the present findings suggest that anhedonic symptoms may specifically impair the emotional processing of pleasant stimuli. Before testing the main hypothesis, the effect of the emotional category was examined. This effect was found to be significant across all ERPs. Specifically, the LPP was larger for both pleasant and unpleasant images compared to neutral ones, indicating heightened elaboration of emotional stimuli. In contrast, the Cue-P300 and SPN were larger for pleasant (but not unpleasant) images relative to neutral ones, reflecting increased affective engagement and anticipation of pleasant stimuli. Most studies have reported increased amplitude of these components during the anticipation of appetitive stimuli using reward tasks [ 40 , 53 , 55 , 57 ]. However, the current findings may extend these previous results, suggesting a general tendency to engage with and anticipate appetitive content, even when the pleasant stimuli are passively presented. Finally, subjective SAM ratings confirmed that emotional pictures were perceived as more arousing than neutral ones, validating the experimental manipulation. Nonetheless, anhedonia did not influence the self-report ratings of valence and arousal. These findings align with other studies suggesting that event-related potentials may be more sensitive than self-report measures in detecting anhedonic symptoms, capturing attentional and emotional processes that are not reflected by self-report ratings [ 49 , 69 ]. This underscores the importance of combining psychophysiological and self-report measures to gain deeper insights into the affective processing patterns underlying vulnerability to depression. Overall, this study contributes at enhancing the understanding of anhedonia as a risk factor for depression. From the RDoC perspective, both clinical depression and its associated risk have been linked to the hypoactivation of the Positive Valence System (PVS), characterized by reduced anticipation and diminished elaboration of appetitive stimuli [ 20 , 48 , 49 , 86 ]. However, the present findings indicate that vulnerability to depression associated with anhedonia may involve a more complex pattern of affective processing, marked by increased anticipation followed by reduced elaboration of pleasant stimuli. Further research may aid in examining this pattern in greater detail, to better delineate anhedonic symptoms within the PVS and across other dimensions of the RDoC framework. From a clinical perspective, these findings provide preliminary insights into the mechanisms underlying anhedonia and may help inform potential approaches to targeted screening and interventions. From these findings, it can be suggested that increasing exposure to rewarding experiences or employing behavioral activation techniques could be particularly beneficial for individuals with anhedonia [ 87 ]. In this context, several clinical interventions have been developed to enhance the activation of the PVS, including Positive Affect Treatment [88], Amplification of Positivity [89], and Behavioral Activation for Anhedonia [ 90 ]. These interventions primarily emphasize the active pursuit of pleasant experiences [ 91 , 92 ] and incorporate exposure to pleasant stimuli, which foster their enhanced elaboration [88,89,90,91,92,93]. Therefore, integrating psychophysiological measures such as ERPs can provide valuable insights into the effectiveness of these interventions. For example, savoring, an emotion regulation technique designed to enhance, prolong, and intensify positive emotions, has been found to significantly increase LPP amplitude during the processing of pleasant stimuli [94,95]. Leveraging ERPs in future research could refine the assessment of depression vulnerability and enhance the specificity of both preventive and clinical interventions. These findings must be considered in light of several limitations. First, the sample was confined to young adults, primarily university students who were not completely free from depressive symptoms [ 65 ]. Second, it was not possible to separate the effects of anhedonia from that of other confounding variables that can impact affective processing, such as anxiety symptoms [ 96 ] having had a depressive episode in the past, or parental history of depression [ 14 , 97 ]. In addition, the SHAPS was the sole measure of anhedonia included in this study [ 63 ]. This scale has good convergent and discriminant validity [98] but only encompasses the construct of consummatory anhedonia [ 11 ]. In sum, this study adds to the current literature exploring how anhedonia, a core symptom and vulnerability factor of depression, impacts neural responses to emotional stimuli at different stages of affective processing. Results indicate that individuals with anhedonia are more likely to experience greater anticipation and reduced elaboration of pleasant stimuli. These findings enhance the understanding of depression vulnerability and may inform the development of targeted preventive interventions aimed at regulating the aberrant processing of pleasant stimuli. Declarations Competing interests The authors declare no competing interests. Author Contribution V.M., C.D.A, and S.M.B. conceived and designed the study; V.M. and R.M. conducted the study; V.M. analyzed the data; V.M., C.D.A., and S.M.B visualized and interpreted the results; V.M. wrote the first draft; all authors reviewed the manuscript. Acknowledgement We acknowledge financial support under the National Recovery and Resilience Plan (NRRP) funded by the European Union – NextGenerationEU – Project Numbers: 20228P4H2K and P20223PTH4, adopted by the Italian Ministry of University and Research (MUR). Data Availability The datasets and materials analyzed in this study are not publicly available due to ethical considerations. They can be requested from the first author and will be provided upon reasonable request. References WHO, Depression and Other Common Mental Disorders: Global Health Estimates, (2017). Demyttenaere, K., et al. Comorbid painful physical symptoms and depression: Prevalence, work loss, and help seeking. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-7073207","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":508665728,"identity":"411d24f7-e67c-4d40-b314-39992b70fe6c","order_by":0,"name":"Valentina Mologni","email":"","orcid":"","institution":"University of Padua","correspondingAuthor":false,"prefix":"","firstName":"Valentina","middleName":"","lastName":"Mologni","suffix":""},{"id":508665729,"identity":"c4787853-5268-4389-989a-624f0ce6798c","order_by":1,"name":"Carola Dell’Acqua","email":"","orcid":"","institution":"University of Padua","correspondingAuthor":false,"prefix":"","firstName":"Carola","middleName":"","lastName":"Dell’Acqua","suffix":""},{"id":508665730,"identity":"72b23be8-2599-4f23-979f-358ae7904c6f","order_by":2,"name":"Roza Mejza","email":"","orcid":"","institution":"University of Padua","correspondingAuthor":false,"prefix":"","firstName":"Roza","middleName":"","lastName":"Mejza","suffix":""},{"id":508665731,"identity":"8c13f1d7-c1a7-40a6-af24-22a28dfaa2a2","order_by":3,"name":"Simone Messerotti Benvenuti","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAABAklEQVRIiWNgGAWjYDACZnQBNvYGBoYEoDgbkVoMJNh4DkC14NSDroVBIgFqFA4t8u3ciY8LKhii+fmPP3xcUfGnjk/yjdmDhzusGfjkG7Cbeph3s/GMMwy5M2fkGBueOQN0mHSOuUHimXScDjNg5t0mzdvGkLvhBg+bZGMbWIuZRGLbYZxa5JtBWv4x5O4/f/z5T7AWyTP4tTAcBmlpANrCkGDGCNYiwYNfC9gvPMckcmfcyDGWbDhjLNnGk1YmAfQLDxtbAnaH9Z/d+Jinxia3v//4w48NFXL88u2Ht0n+3GEtJ998AIfLwEACjc/YwMCDTz0WANQyCkbBKBgFowAGAEHsT1N9Fa2YAAAAAElFTkSuQmCC","orcid":"","institution":"University of Padua","correspondingAuthor":true,"prefix":"","firstName":"Simone","middleName":"Messerotti","lastName":"Benvenuti","suffix":""}],"badges":[],"createdAt":"2025-07-08 09:38:10","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-7073207/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-7073207/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":90464633,"identity":"d687045c-bb4a-48bc-aa52-8db8be3e0af9","added_by":"auto","created_at":"2025-09-03 05:10:02","extension":"jpeg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":131797,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003e(Panel a) \u003c/strong\u003eGrand average ERP waveforms for a full epoch trial of the S1-S2 task at parietal electrodes (P3, PZ, P4). Cue (S1) onset was at 0 s, followed by the image (S2) onset at 6 s. \u003cstrong\u003e(Panel b)\u003c/strong\u003e Grand average of the Cue-P300, scored as the peak amplitude in the shaded window (0.2 – 0.4 s). \u003cstrong\u003e(Panel c)\u003c/strong\u003e Grand average of the LPP, scored as the mean amplitude in the shaded window (6.5 to 7 s). \u003cstrong\u003e(Panel d)\u003c/strong\u003e Topographic maps of the LPP for unpleasant, neutral, and pleasant images.\u003c/p\u003e","description":"","filename":"Figure107072025.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-7073207/v1/58dc7dd27749f8267257c05c.jpeg"},{"id":90464637,"identity":"6b1fce31-b336-4a4d-b096-d3c203bf51fc","added_by":"auto","created_at":"2025-09-03 05:10:02","extension":"jpeg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":118888,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003e(Panel a) \u003c/strong\u003eGrand average ERP waveforms during the S1-S2 task at frontocentral electrodes (FC1, FC2, FC5, FC6). \u003cstrong\u003e(Panel b) \u003c/strong\u003eGrand average of the SPN, scored as the mean amplitude in the shaded window (5.8 – 6 s). \u003cstrong\u003e(Panel c)\u003c/strong\u003e Topographic maps of the SPN for unpleasant, neutral, and pleasant images.\u003c/p\u003e","description":"","filename":"Figure207072025.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-7073207/v1/17769d9903cf07f6b83b5cb4.jpeg"},{"id":90464639,"identity":"12abc821-3c61-498e-9ddf-24e487f8009e","added_by":"auto","created_at":"2025-09-03 05:10:02","extension":"jpeg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":124325,"visible":true,"origin":"","legend":"\u003cp\u003eInteraction effect of SHAPS (mean-centered values) and emotional category (pleasant, unpleasant) on ΔSPN amplitude. Ninety-five % confidence bands are presented in different colors.\u003c/p\u003e","description":"","filename":"Figure307072025.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-7073207/v1/9bb9d4a9af19946b2bae3553.jpeg"},{"id":90466738,"identity":"55610a39-4889-4146-af45-f90993b33c9f","added_by":"auto","created_at":"2025-09-03 05:36:46","extension":"jpeg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":118474,"visible":true,"origin":"","legend":"\u003cp\u003eInteraction effect of SHAPS (mean-centered values) and emotional category (pleasant, unpleasant) on ΔLPP scores. Ninety-five % confidence bands are presented in different colors.\u003c/p\u003e","description":"","filename":"Figure407072025.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-7073207/v1/6b07f2eeba80c181a6570871.jpeg"},{"id":92696854,"identity":"d502d051-393f-4af9-b88b-bc2cf351c72e","added_by":"auto","created_at":"2025-10-03 07:09:35","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1372784,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7073207/v1/8f4c1f4f-f87d-4228-9f9c-aaecae4d4b17.pdf"},{"id":90464634,"identity":"96379200-fc7c-4bec-9da2-c81ad6ee39e2","added_by":"auto","created_at":"2025-09-03 05:10:02","extension":"docx","order_by":0,"title":"","display":"","copyAsset":false,"role":"supplement","size":1931298,"visible":true,"origin":"","legend":"","description":"","filename":"SuppAnhedERPs07072025.docx","url":"https://assets-eu.researchsquare.com/files/rs-7073207/v1/598a06113aa4d2b32109c51d.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"The time course of affective processing in anhedonia: insights from event-related potentials","fulltext":[{"header":"Introduction","content":"\u003cp\u003eDepression is recognized as a leading cause of disability, affecting over 300\u0026nbsp;million people globally [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e]. It manifests through a range of affective, cognitive, and somatic symptoms [\u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e]. Given its substantial burden, identifying viable factors contributing to depression vulnerability has been highlighted as a core priority [\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e]. In this regard, emerging research suggests that anhedonia—defined as the loss of pleasure and emotional disengagement from appetitive stimuli—may play a central role in the disorder's onset and maintenance, being a potential endophenotype for depression [\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e]. Notably, anhedonia can precede the onset of depression and often persists even after other symptoms have remitted [\u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e]. However, anhedonia is not a unitary construct, and addressing its heterogeneity could enhance our understanding of depression vulnerability [\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e7\u003c/span\u003e].\u003c/p\u003e\u003cp\u003eAligned with this purpose, the Research Domain Criteria (RDoC) initiative proposed a dimensional model whereby integrating multiple levels of analysis of different constructs might be useful to fully characterize anhedonia. Among the RDoC constructs, the Positive Valence Systems (PVS) may be particularly relevant for its understanding, as they encompass a set of systems involved in anticipating, obtaining, elaborating, and responding to pleasant or rewarding stimuli [\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e, \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e]. While these phases are related, they are also dissociable at both behavioral and neural levels, each of them serving distinct functions: anticipation allows individuals to prepare for future events, while elaboration and response mobilization entail engaging with and processing the immediate reward. Reduced appetitive anticipation and elaboration has been observed in depression [\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e, \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e], in children with familial risk of depression [\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e], and it can prospectively predict its onset [\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e, \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e, \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e]. Yet, it is still unclear whether depression vulnerability stems from deficits occurring at any specific stage of appetitive stimulus processing [\u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e, \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e]. For example, anhedonia may not necessarily reflect a diminished ability to experience pleasure, but rather a deficit in mobilizing motivational resources to pursue pleasurable activities [\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e].\u003c/p\u003e\u003cp\u003eStudies employing functional magnetic resonance imaging (fMRI) helped disentangle the existence of both shared and dissociable neural mechanisms related to distinct phases of appetitive stimulus processing [\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e]. Specifically, the anticipation of appetitive stimuli consistently engages the ventral striatum, while the elaboration phase is related with heightened activity in the orbitofrontal cortex [\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e, \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e, \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e] and regulatory involvement of dorsolateral and ventromedial prefrontal cortices [\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e26\u003c/span\u003e]. Notably, altered cortical and striatal activation during the anticipation and processing of appetitive stimuli has been observed in adults with current depression [\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e, \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e, \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e], in remission [\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e], and in youths at risk for depression [\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e]. However, most of these studies have relied upon monetary reward tasks, without examining the distinct phases of other appetitive stimuli processing (e.g., pleasant images, social rewards) that might be more relevant to the anhedonic symptomatology in the context of depression risk [\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e, \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e]. For example, greater neural responses to pleasant pictures but not monetary rewards have been associated with improved treatment outcomes for depression [\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e]. More importantly, due to its poor temporal resolution, fMRI might be inadequate for the online tracking of the distinct phases of appetitive stimulus processing, leading to the merging of neural activity associated with processes that are temporally close but psychologically distinct (i.e., anticipation, stimulus response, elaboration) [\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e].\u003c/p\u003e\u003cp\u003eThese issues can be addressed by using event-related potentials (ERPs) of the electroencephalographic (EEG) signal [\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e]. Thanks to their excellent temporal resolution, ERPs allow for the precise examination of distinct phases of appetitive stimulus processing. An extensively used paradigm in ERP research to tackle the anticipation and elaboration of affective stimuli is the S1-S2 task, which involves the presentation of a series of paired trials in which an initial cue stimulus (S1) indicates the emotional valence of a second stimulus (S2, or imperative stimulus) that appears after a fixed inter-stimulus interval (ISI) [\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e37\u003c/span\u003e, \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e, \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e39\u003c/span\u003e]. This allows the dissociation of anticipation from the elaboration of appetitive stimuli in the brain. The Cue-P300 and the Stimulus Preceding Negativity (SPN) are two main components associated with anticipation, whereas elaboration is associated with the Late Positive Potential (LPP). The Cue-P300 is a positive parietal deflection occurring at about 300 ms after cue (S1) onset, which reflects affective engagement (i.e., the allocation of attentional resources engaged in the upcoming stimulus) towards the upcoming stimulus (S2) and is typically larger for emotional vs. neutral stimuli [\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e]. In addition, the Stimulus Preceding Negativity (SPN) is a slow negative potential that emerges approximately 200 ms before the S2, exhibiting a larger amplitude (i.e., a greater negativity) during the anticipation of emotional vs. neutral stimuli at frontocentral sites [\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e, \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e42\u003c/span\u003e]. The SPN is linked to the anticipation of emotionally arousing content [\u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e43\u003c/span\u003e] and is modulated by the level of motivational engagement with the forthcoming stimulus. Finally, the LPP is a positive waveform that begins approximately 400 ms following the S2 onset, with maximal amplitude at centro-parietal sites. This ERP reflects elaborative processing, including motivated attention, as well as stimulus representation and evaluation [\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e44\u003c/span\u003e]. In addition, emotional stimuli consistently elicit a larger LPP compared to neutral stimuli [45] reflecting the activation of the appetitive motivational system in the brain [\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e46\u003c/span\u003e].\u003c/p\u003e\u003cp\u003eInvestigating whether individuals who are vulnerable to depression – such as those experiencing anhedonic symptoms – exhibit affective processing alterations which are analogous to those observed in individuals with clinical depression may help identify early indicators of the disorder and clarify whether these alterations are potential precursors rather than mere characteristics of the clinical condition. Consistently, research on affective processing has reported a blunted LPP to pleasant stimuli in not only in individuals with diagnosed depression [\u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e47\u003c/span\u003e, \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e48\u003c/span\u003e], but also in those at risk for the disorder, such as individuals with subclinical symptoms or familial risk for depression [49,50,51]. This finding suggests that reduced affective elaboration of pleasant content, indexed by a blunted LPP amplitude, may serve as a psychophysiological marker of depression vulnerability, potentially reflecting anhedonia [\u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e48\u003c/span\u003e]. In line with this, the limited number of studies examining the two ERP components related to anticipation (i.e., Cue-P300 and SPN) have found a reduction in the Cue-P300 amplitude in response to rewarding stimuli among individuals with both subclinical and clinical depression [\u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e52\u003c/span\u003e, \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e53\u003c/span\u003e]. Interestingly, blunted Cue-P300 has been observed in individuals with anhedonia but without current depressive symptoms during a gambling task [\u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e54\u003c/span\u003e], suggesting that anhedonia may influence affective engagement for future rewards. In contrast, studies using reward tasks have yielded mixed findings regarding the SPN, likely due to differences in task design and that this component is sensitive to several manipulations [38, 55, 56, 57]. For instance, the SPN amplitude has been reported as blunted [\u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e58\u003c/span\u003e], increased [59], or unchanged [\u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e52\u003c/span\u003e] in individuals with depressive symptoms relative to controls. Moreover, two studies using reward tasks and assessing anhedonia while controlling for depressive symptoms reported no significant change in SPN amplitude during the anticipation of rewarding relative to non-rewarding stimuli [\u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e54\u003c/span\u003e, \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e60\u003c/span\u003e]. Given that most of the ERPs research has relied on traditional reward-based paradigms — primarily assessing the processing of external incentives such as monetary rewards — the S1–S2 task enables the investigation of intrinsic affective responses to emotionally salient stimuli, such as pleasant images [\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e37\u003c/span\u003e, \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e]. This distinction is particularly important, as heightened neural responses to pleasant images – but not to monetary rewards – have been associated with more favorable treatment outcomes in individuals with depression [\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e]. In addition, the temporal dynamics of affective processing of pleasant stimuli remain underexplored, as most existing studies have focused exclusively on the elaboration stage [\u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e47\u003c/span\u003e, \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e48\u003c/span\u003e, \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e49\u003c/span\u003e, \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e50\u003c/span\u003e].\u003c/p\u003e\u003cp\u003eThe reviewed literature suggests that appetitive stimuli processing might contribute to anhedonia and, consequently, depression vulnerability. However, there is still limited evidence examining the link between anhedonia and the distinct appetitive picture processing stages. Hence, this study aimed to examine appetitive picture processing through the assessment of ERPs during an emotional S1-S2 task in a sample of young adults with varying levels of anhedonia. Based on the reviewed literature, it was hypothesized that higher levels of anhedonia would be associated with reduced affective elaboration of pleasant stimuli, as indicated by a blunted LPP amplitude. Moreover, although the literature is still scarce, it was hypothesized that higher levels of anhedonia would be associated with reduced appetitive cue engagement and anticipation, as indexed by decreased amplitude of Cue-P300 and SPN components. While the primary focus was on appetitive processing in anhedonia, an exploratory analysis of the affective processing of unpleasant stimuli was also conducted. Lastly, this study explored whether the observed appetitive processing dynamics are uniquely associated with anhedonia or extend to non-anhedonic depressive symptoms.\u003c/p\u003e"},{"header":"Methods","content":"\u003cp\u003e\u003cem\u003eParticipants\u003c/em\u003e\u003c/p\u003e\u003cp\u003eForty-five Italian Caucasian Students (31 females, mean (M) age = 21.7 years, standard deviation (SD) = 1.84 years, range = 19–26 years) from the University of Padua (Italy) voluntarily participated in the study. An a-priori power analysis was performed using G*Power 3 [\u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e61\u003c/span\u003e]. According to methodological guidelines suggesting that ERPs are adequately powered to detect large effects [\u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e62\u003c/span\u003e], the power analysis indicated that a sample size of 41 participants would be sufficient to detect a medium-to-large effect size (f = 0.3) with a statistical power of 0.8 (α = 0.05, predictors = 3). The enrolled sample was medically healthy and free from psychotropic medication, as assessed with an ad-hoc interview. Exclusion criteria included a current diagnosis or history of cardiovascular and neurological diseases and a formal diagnosis of mental diseases. All participants had a normal-to-corrected vision and were naïve to the purpose of the experiment. All participants understood and signed the informed consent forms. The study was conducted in compliance with the Declaration of Helsinki on research on human subjects and was approved by the Ethical Committee of Psychological Research, Area 17, University of Padua (prot. no. 564-b). Participants received a monetary compensation of 25 euros for their participation.\u003c/p\u003e\u003cp\u003e\u003cem\u003ePsychological measures\u003c/em\u003e\u003c/p\u003e\u003cp\u003eThe Snaith-Hamilton Pleasure Scale (SHAPS) was used to assess anhedonia (internal consistency: Kuder-Richardson coefficient = 0.85) [\u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e63\u003c/span\u003e]. The SHAPS consists of 14 items that cover four domains of pleasure experience: interest, social interaction, sensory experience, and food/drink. Each item (e.g. I would be able to enjoy a beautiful landscape or view) is rated on a 4-point scale from “strongly disagree” to “strongly agree.” Responses of “disagree” score 1 point each, while “agree” responses score 0 point each. The total score ranges from 0 to 14, with scores of 3 or higher indicating a reduced ability to experience pleasure, and consequently, the presence of anhedonic symptoms.\u003c/p\u003e\u003cp\u003eTo evaluate depressive symptoms over the past two weeks, the Italian version of the Beck Depression Inventory-II [\u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e64\u003c/span\u003e, \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e65\u003c/span\u003e] was administered. This self-report questionnaire comprises 21 items, each rated on a four-point Likert scale, with total scores ranging from 0 to 63 (internal consistency: Cronbach's alpha = 0.87). Higher scores indicate more severe depressive symptoms. In the Italian version, a score of 12 is the cut-off point for distinguishing between individuals with and without depressive symptoms [\u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e65\u003c/span\u003e]. From the BDI-II, a non-anhedonic symptom (sum of all items except 4, 12, 22) subscale was derived (internal consistency: Cronbach's alpha = 0.83) [\u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e66\u003c/span\u003e].\u003c/p\u003e\u003cp\u003e\u003cem\u003eExperimental task\u003c/em\u003e\u003c/p\u003e\u003cp\u003e In this study, participants engaged in an S1-S2 emotional paradigm while undergoing electroencephalographic (EEG) recording. The task consisted of a single block of 84 trials. Each trial started with a 500 ms baseline (a white fixation dot), followed by a cue (S1) lasting 1500 ms that signaled the emotional content (a plus for pleasant, a circle for neutral, a minus for unpleasant) of the upcoming picture (S2), presented after the 4500 ms inter-stimulus interval (ISI) and lasting 6000 ms. The S2 was followed by a variable inter-trial interval (ITI) of 6000–8000 ms, during which a white fixation dot (identical to the baseline) was presented (see Supplementary, Figure \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e online). Participants were instructed to observe the cue (S1) and the subsequent emotional image (S2), with no motor response required. They were also informed that the emotional image would always be preceded by a congruent cue, which would anticipate its emotional valence. The S2 stimuli comprised 42 emotional images (600 x 800 pixels), each repeated twice, selected from the International Affective Picture System [\u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e67\u003c/span\u003e] including 14 pleasant images (i.e., erotic couples, sports), 14 neutral images (i.e., neutral scenes, household objects), and 14 unpleasant images (i.e., attacking humans, explosions, animals). The selection was based on normative arousal ratings, ensuring that pleasant and unpleasant images were matched in arousal levels (pleasant, mean ± SD = 6.5 ± 0.3; unpleasant, mean ± SD = 6.2 ± 0.6; \u003cem\u003ep\u003c/em\u003e = .11) which were significantly higher than those of neutral pictures (neutral, mean ± SD = 3.1 ± 0.5, all \u003cem\u003eps\u003c/em\u003e \u0026lt; .001) [\u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e68\u003c/span\u003e]. To induce greater psychophysiological changes, only highly arousing pleasant and unpleasant pictures were selected [\u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e69\u003c/span\u003e]. Stimuli were presented in a semi-randomized sequence to prevent consecutive S1-S2 pairs from having the same emotional valence. At the end of the task, the 42 images (14 for each emotional category) shown during the S1-S2 paradigm were presented again, and ratings of emotional valence and arousal were obtained using a computerized version of the 9-point Valence and Arousal scales of the Self-Assessment Manikin (SAM) [70]. (IAPS catalogue numbers: 1120, 1201, 1301, 1525, 2493, 2495, 2512, 2575, 2595, 2840, 4611, 4647, 4651, 4652, 4670, 4695, 5532, 6200, 6250, 6260, 6312, 7002, 7004, 7009, 7036, 7041, 7224, 7547, 8034, 8080, 8180, 8185, 8200, 8370, 8400, 8490, 9185, 9622, 9630, 9635.2, 9909, 9940).\u003c/p\u003e\u003cp\u003e\u003cem\u003eProcedure\u003c/em\u003e\u003c/p\u003e\u003cp\u003eThe study was advertised via various means, including flyers and online platforms. Interested students from the University of Padua completed an online form, administered through Google Forms. The form encompassed an anamnestic interview to assess eligibility, the SHAPS, and the BDI-II. Eligible participants were invited to a laboratory session in the Clinical Psychophysiology laboratory at the Department of General Psychology of the University of Padua. Participants were instructed to refrain from alcohol consumption on the day before the experimental session and to avoid caffeine and nicotine for at least four hours before the session. Upon arrival at the laboratory, participants read and signed the informed consent form, after which they underwent an ad-hoc anamnestic interview. Participants were then seated on a comfortable chair in a dimly lit, sound-attenuated room. This study was part of a broader project that also included the recording of the electrocardiogram (ECG), in addition to the EEG. After electrode attachment and a 3-minute resting-state recording, they were given three practice trials, one for each emotional category (pleasant, neutral, unpleasant). Following the practice trials, participants completed the S1-S2 paradigm. The entire procedure took approximately 90 min.\u003c/p\u003e\u003cp\u003e\u003cem\u003eElectroencephalogram data acquisition and analysis\u003c/em\u003e\u003c/p\u003e\u003cp\u003eEEG was recorded using a 32-channel ANT system and a computer running eego™ software (ANT Neuro, Enschede, Netherlands). The elastic cap with 32 tin electrodes was arranged according to the 10–20 System (FP1, FPz, FP2, F7, F3, Fz, F4, F8, FC5, FC1, FC2, FC6, T7, C3, Cz, C4, T8, CP5, CP1, CP2, CP6, P7, P3, Pz, P4, P8, POz, O1, Oz, O2, and M1 and M2 [mastoids]), referenced online to CPz. Vertical and horizontal electrooculograms (EOGs) were recorded using a bipolar montage to track horizontal eye movements and blinks, with electrodes placed above and below the right orbit and at the external canthi of the eyes. Electrode impedance was kept below 10 kΩ. The EEG and the EOG signals were recorded in DC with a low-pass filter of 30 Hz and sampled at 1000 Hz. EEG data was resampled to 500 Hz and re-referenced offline to a linked mastoids montage in EEGLAB [\u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e71\u003c/span\u003e]. Further analyses were performed in Brainstorm [\u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e72\u003c/span\u003e]. Data was band-pass filtered from 0.01 to 30 Hz and corrected for blink artifacts using Independent Component Analysis (ICA). The EEG signal was segmented into 8500 ms epochs, from 500 ms before S1 to 2000 ms after the onset of S2 (i.e., -500 to 8000 ms). Based on previous research, the signal was baseline-corrected using the average signal activity within the − 250 ms to -50 ms time window preceding S1. This window was selected for each ERP, as it was the period of EEG recording being closest to the participant's true baseline, without other cognitive or emotional processes occurring [\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e, \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e73\u003c/span\u003e]. A semiautomatic procedure was employed for artifact rejection, applying a voltage difference threshold of 200 µV (peak-to-peak). Then, epochs were visually inspected to detect and reject possible remaining artifacts. According to previous research [38,74,75] and the visual inspection of the grand-average ERPs waveforms for the three emotional categories, the Cue-P300 and the LPP were maximal at parietal sites (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e), while the SPN was prominent at frontocentral sites (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Therefore, the Cue-P300 was calculated by averaging the peak amplitudes at the parietal poll (P3, PZ, P4) in the time window from 200 to 400 ms post-S1, while the SPN was calculated as the mean amplitude at the frontocentral poll (FC1, FC2, FC5, FC6) within the 200 ms preceding the onset of S2. Further, the LPP was calculated by averaging the mean amplitude at the parietal poll (P3, PZ, P4) in the time window from 500 to 1000 ms post-S2. The grand average ERP waveforms for parietal, central, and frontal electrodes during the S1-S2 task, are presented in the supplementary material (see Supplementary, Figure S2 online).\u003c/p\u003e\u003cp\u003e\u003cem\u003eStatistical Analyses\u003c/em\u003e\u003c/p\u003e\u003cp\u003eStatistical analyses were conducted in Rstudio [\u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e76\u003c/span\u003e]. A \u003cem\u003ep\u003c/em\u003e-value of .05 was used as the threshold for statistical significance. Experimental hypotheses were tested using linear mixed-effect models with the \u003cem\u003elme4\u003c/em\u003e and \u003cem\u003elmerTest\u003c/em\u003e packages [\u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e77\u003c/span\u003e, \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e78\u003c/span\u003e]. All models included participants as a random intercept, and sex (biological) as a covariate to account for the sex unbalance within the experimental sample.\u003c/p\u003e\u003cp\u003eTo test the effect of emotional category and anhedonia on self-reported ratings of valence and arousal, two separate models with Category (i.e., pleasant, neutral, unpleasant), SHAPS scores, and their interaction as fixed factors were conducted: Model \u003cem\u003e← lmer(Subjective rating of arousal or valence ∼ Category × SHAPS + Sex + (1|Subject)).\u003c/em\u003e\u003c/p\u003e\u003cp\u003eThen, to ensure the effect of emotional valence on the Cue-P300, SPN, and LPP, three separate models with Category as a fixed factor were conducted on each ERP: \u003cem\u003eModel ← lmer(ERP amplitude ∼ Category + Sex + (1|Subject)).\u003c/em\u003e\u003c/p\u003e\u003cp\u003eTo test the primary hypothesis—examining the relationship between anhedonia levels and distinct stages of emotional processing—three separate models were conducted, one for each ERP component (Cue-P300, SPN, LPP). Differential ERP scores (ΔERP; pleasant - neutral, unpleasant - neutral) were employed to reduce the number of predictors and isolate the effect of the emotional category [\u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e79\u003c/span\u003e]. The models included Category (pleasant, unpleasant), SHAPS scores, and their interaction as fixed factors: \u003cem\u003eModel ← lmer(△ERPs amplitude ∼ Category × SHAPS + Sex + (1|Subject)).\u003c/em\u003e To test whether the examined effects are uniquely related to anhedonic symptoms, the same models were conducted with non-anhedonic depressive symptoms as predictor. Particularly, Category (pleasant, unpleasant), BDI-II non-anhedonic scores, and their interaction were included as fixed effects in the model: \u003cem\u003eModel ← lmer(△ERPs amplitude ∼ Category × BDI-II non-anhedonic + Sex + (1|Subject)).\u003c/em\u003e\u003c/p\u003e\u003cp\u003eFor the fixed effects, the estimated coefficients (\u003cem\u003eb\u003c/em\u003e), standard errors (SE), \u003cem\u003et\u003c/em\u003e values, and confidence intervals for each parameter were reported. Additionally, \u003cem\u003ep\u003c/em\u003e-values obtained through the Satterthwaite approximation, as implemented in the \u003cem\u003elmerTest\u003c/em\u003e package [\u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e78\u003c/span\u003e], were provided. All predictors were centered and scaled. The collinearity was tested by calculating the Variance Inflation Factors (VIF) with the \u003cem\u003evif\u003c/em\u003e function of the car package [\u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e80\u003c/span\u003e]. Significant categorical effects (\u003cem\u003ep\u003c/em\u003e \u0026lt; .05) were followed by Tukey HSD post-hoc tests to correct for multiple comparisons.\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003e\u003cem\u003ePsychological measures and valence and arousal self-report ratings\u003c/em\u003e\u003c/p\u003e\u003cp\u003eThe average SHAPS score was 1.8 (SD\u0026thinsp;=\u0026thinsp;1.7, range\u0026thinsp;=\u0026thinsp;0\u0026ndash;6), with 16 participants (35.55% of the total sample) scoring above the cut-off (i.e., \u0026ge; 3). The average BDI-II score was 13.82 (SD\u0026thinsp;=\u0026thinsp;10.11, range\u0026thinsp;=\u0026thinsp;1\u0026ndash;45), with 24 participants (53.33 % of the total sample) scoring above the cut off (i.e., \u0026ge; 12). Additionally, the average BDI-II non-anhedonic score was 12.22 (SD\u0026thinsp;=\u0026thinsp;8.86, range = \u0026minus;\u0026thinsp;39). Linear mixed-effects models on self-reported SAM ratings revealed a significant main effect of Category in predicting valence (F\u003csub\u003e(2,1843)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;907.15, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;.001) and arousal (F\u003csub\u003e(2,1843)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;517.12, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;.001). Specifically, subjective ratings of valence were significantly higher for pleasant images compared to neutral and unpleasant images (all \u003cem\u003eps\u003c/em\u003e\u003csub\u003e\u003cem\u003eTukey\u003c/em\u003e\u003c/sub\u003e \u0026lt; .001), and significantly lower for unpleasant images compared to neutral images (\u003cem\u003ep\u003c/em\u003e\u003csub\u003e\u003cem\u003eTukey\u003c/em\u003e\u003c/sub\u003e \u0026lt; .001). In addition, subjective ratings of arousal were significantly higher for both pleasant and unpleasant compared to neutral images (all \u003cem\u003eps\u003c/em\u003e\u003csub\u003e\u003cem\u003eTukey\u003c/em\u003e\u003c/sub\u003e \u0026lt; .001), but also for unpleasant compared to pleasant images (\u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;.002). No significant effect of SHAPS or Category \u0026times; SHAPS (all \u003cem\u003eps\u003c/em\u003e\u0026thinsp;\u0026gt;\u0026thinsp;.09) emerged. Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e shows the means and standard deviations of SAM valence and arousal ratings.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eMean and standard deviation of valence and arousal ratings for pleasant, neutral, and unpleasant pictures in the current sample.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"4\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSAM ratings\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003ePleasant\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eNeutral\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eUnpleasant\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eValence\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c2\"\u003e\u003cp\u003e6.78\u0026thinsp;\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e\u0026plusmn;\u003c/span\u003e\u0026thinsp;1.18\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e\u003cp\u003e5.20\u0026thinsp;\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e\u0026plusmn;\u003c/span\u003e\u0026thinsp;1.30\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e\u003cp\u003e3.17\u0026thinsp;\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e\u0026plusmn;\u003c/span\u003e\u0026thinsp;1.65\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eArousal\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c2\"\u003e\u003cp\u003e4.94\u0026thinsp;\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e\u0026plusmn;\u003c/span\u003e\u0026thinsp;2.37\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e\u003cp\u003e2.47\u0026thinsp;\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e\u0026plusmn;\u003c/span\u003e\u0026thinsp;2.36\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e\u003cp\u003e5.26\u0026thinsp;\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e\u0026plusmn;\u003c/span\u003e\u0026thinsp;1.68\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003ctfoot\u003e\u003ctr\u003e\u003ctd colspan=\"4\"\u003e\u003cem\u003eNote\u003c/em\u003e. Data are M\u0026thinsp;\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e\u0026plusmn;\u003c/span\u003e\u0026thinsp;SD; SAM\u0026thinsp;=\u0026thinsp;Self-Assessment Manikin.\u003c/td\u003e\u003c/tr\u003e\u003c/tfoot\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003cem\u003eThe influence of emotional category on ERPs amplitude\u003c/em\u003e\u003c/p\u003e\u003cp\u003eThe mixed-effect models predicting the ERPs amplitude from the emotional Category revealed a significant effect of Category across all the ERPs (Cue-P300, F\u003csub\u003e(2,358)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;10.27, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;.01; SPN, F\u003csub\u003e(2,487)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;3.18, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;=\u0026thinsp;.04; LPP, F\u003csub\u003e(2,358)\u003c/sub\u003e\u0026thinsp;=\u0026thinsp;21.17, \u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;.01). Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e shows the mean ERPs amplitude for each emotional Category. In the Cue-P300 model, the peak amplitude was larger for pleasant relative to neutral and unpleasant trials (all \u003cem\u003eps\u003c/em\u003e\u003csub\u003e\u003cem\u003eTukey\u003c/em\u003e\u003c/sub\u003e \u003cem\u003e\u0026lt;\u003c/em\u003e .01), while no significant difference emerged between unpleasant and neutral trials (\u003cem\u003ep\u003c/em\u003e\u003csub\u003e\u003cem\u003eTukey\u003c/em\u003e\u003c/sub\u003e = .21). In the SPN model, the mean amplitude was larger (i.e., more negative) for pleasant relative to neutral trials (\u003cem\u003ep\u003c/em\u003e\u003csub\u003e\u003cem\u003eTukey\u003c/em\u003e\u003c/sub\u003e = .03), while no significant difference emerged between unpleasant relative to pleasant or unpleasant relative to neutral trials (all \u003cem\u003eps\u003c/em\u003e\u003csub\u003e\u003cem\u003eTukey\u003c/em\u003e\u003c/sub\u003e \u003cem\u003e\u0026gt;\u003c/em\u003e .24). Finally, the LPP mean amplitude was larger for emotional (i.e., pleasant, unpleasant) relative to neutral trials (all \u003cem\u003eps\u003c/em\u003e\u003csub\u003e\u003cem\u003eTukey\u003c/em\u003e\u003c/sub\u003e \u003cem\u003e\u0026lt;\u003c/em\u003e .01), while no significant difference between pleasant and unpleasant trials emerged (\u003cem\u003ep\u003c/em\u003e\u003csub\u003e\u003cem\u003eTukey\u003c/em\u003e\u003c/sub\u003e = .18). Figure\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e shows the grand averages of the Cue-P300 and LPP, while Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e shows the grand average of the SPN for the three emotional categories.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eMean and standard deviation of the ERPs amplitude (\u0026micro;V) across emotional Categories.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"4\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"char\" char=\"\u0026plusmn;\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colspan=\"4\" nameend=\"c4\" namest=\"c1\"\u003e\u003cp\u003eERPs amplitude (\u0026micro;V)\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eCue-P300\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003eSPN\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003eLPP\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cem\u003eCategory\u003c/em\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePleasant\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c2\"\u003e\u003cp\u003e9.74\u0026thinsp;\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e\u0026plusmn;\u003c/span\u003e\u0026thinsp;4.06\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e\u003cp\u003e1.79\u0026thinsp;\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e\u0026plusmn;\u003c/span\u003e\u0026thinsp;5.15\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e\u003cp\u003e9.92\u0026thinsp;\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e\u0026plusmn;\u003c/span\u003e\u0026thinsp;7.09\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eNeutral\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c2\"\u003e\u003cp\u003e8.59\u0026thinsp;\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e\u0026plusmn;\u003c/span\u003e\u0026thinsp;3.50\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e\u003cp\u003e3.08\u0026thinsp;\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e\u0026plusmn;\u003c/span\u003e\u0026thinsp;6.62\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e\u003cp\u003e7.30\u0026thinsp;\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e\u0026plusmn;\u003c/span\u003e\u0026thinsp;6.79\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eUnpleasant\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c2\"\u003e\u003cp\u003e9.02\u0026thinsp;\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e\u0026plusmn;\u003c/span\u003e\u0026thinsp;3.84\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c3\"\u003e\u003cp\u003e2.62\u0026thinsp;\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e\u0026plusmn;\u003c/span\u003e\u0026thinsp;5.95\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"char\" char=\"\u0026plusmn;\" colname=\"c4\"\u003e\u003cp\u003e10.94\u0026thinsp;\u003cspan type=\"Underline\" class=\"Underline\" name=\"Emphasis\"\u003e\u0026plusmn;\u003c/span\u003e\u0026thinsp;8.97\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003ctfoot\u003e\u003ctr\u003e\u003ctd colspan=\"4\"\u003e\u003cem\u003eNote\u003c/em\u003e. Data are M (SD).\u003c/td\u003e\u003c/tr\u003e\u003c/tfoot\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003e\u003cem\u003eAnhedonia and the distinct phases of appetitive picture processing\u003c/em\u003e\u003c/p\u003e\u003cp\u003eTable\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e shows the results of the mixed-effect model predicting ERPs amplitude from Category, SHAPS, and their interaction. VIF values were all \u0026lt;\u0026thinsp;1.22, indicating acceptable levels of multicollinearity among the predictor variables.\u003c/p\u003e\u003cp\u003e\u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e\u003ccaption language=\"En\"\u003e\u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e\u003cdiv class=\"CaptionContent\"\u003e\u003cp\u003eLinear mixed-models predicting △ERPs amplitudes from emotional category (pleasant, unpleasant) and SHAPS scores.\u003c/p\u003e\u003c/div\u003e\u003c/caption\u003e\u003ccolgroup cols=\"4\"\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e\u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e\u003cthead\u003e\u003ctr\u003e\u003cth align=\"left\" colname=\"c1\"\u003e\u003cp\u003ePredictor\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c2\"\u003e\u003cp\u003eb (SE)\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cem\u003et\u003c/em\u003e\u003c/p\u003e\u003c/th\u003e\u003cth align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cem\u003ep\u003c/em\u003e\u003c/p\u003e\u003c/th\u003e\u003c/tr\u003e\u003c/thead\u003e\u003ctbody\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"4\" nameend=\"c4\" namest=\"c1\"\u003e\u003cp\u003e\u003cb\u003eCue-P300 model\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eCategory\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003e0.72 (0.25)\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cb\u003e2.87\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;.01***\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSHAPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.60 (0.39)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e1.51\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e.13\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSex\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.70 (0.81)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.86\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e.39\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCategory \u0026times;\u0026thinsp;SHAPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.17 (0.25)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.68\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e.49\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"4\" nameend=\"c4\" namest=\"c1\"\u003e\u003cp\u003e\u003cb\u003eSPN model\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCategory\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-0.84 (0.57)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-1.47\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e.14\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSHAPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.13 (0.91)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.14\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e.89\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSex\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.19 (1.88)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.10\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e.91\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eCategory \u0026times;\u0026thinsp;SHAPS\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003e-1.62 (0.58)\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cb\u003e-2.80\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;.01***\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colspan=\"4\" nameend=\"c4\" namest=\"c1\"\u003e\u003cp\u003e\u003cb\u003eLPP model\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eCategory\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-1.02 (0.64)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-1.60\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e.11\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSHAPS\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e0.56 (0.81)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e0.69\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e.49\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003eSex\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e-1.26 (1.60)\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e-0.78\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e.43\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003ctr\u003e\u003ctd align=\"left\" colname=\"c1\"\u003e\u003cp\u003e\u003cb\u003eCategory \u0026times;\u0026thinsp;SHAPS\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c2\"\u003e\u003cp\u003e\u003cb\u003e-1.99 (0.64)\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c3\"\u003e\u003cp\u003e\u003cb\u003e-3.11\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003ctd align=\"left\" colname=\"c4\"\u003e\u003cp\u003e\u003cb\u003e\u0026lt;\u0026thinsp;.01***\u003c/b\u003e\u003c/p\u003e\u003c/td\u003e\u003c/tr\u003e\u003c/tbody\u003e\u003c/colgroup\u003e\u003ctfoot\u003e\u003ctr\u003e\u003ctd colspan=\"4\"\u003e\u003cem\u003eNotes.\u003c/em\u003e The reference levels are Category-unpleasant and Sex-females. Significant results are shown in bold, with asterisks indicating the level of statistical significance (*\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;.05, **\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;.01, ***\u003cem\u003ep\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;.001).\u003c/td\u003e\u003c/tr\u003e\u003c/tfoot\u003e\u003c/table\u003e\u003c/div\u003e\u003c/p\u003e\u003cp\u003eThe model predicting Cue-P300 peak amplitude showed an effect of Category (i.e., the mean amplitude was larger for pleasant vs. unpleasant images), but no other significant effect emerged. The model predicting the SPN amplitude revealed a significant interaction between Category and SHAPS, showing that higher levels of anhedonia were associated with increased (i.e., more negative) SPN to pleasant stimuli compared to unpleasant ones (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). In addition, the model predicting LPP amplitude revealed a significant interaction between Category and SHAPS, indicating that higher levels of anhedonia were associated with reduced LPP to pleasant stimuli compared to unpleasant ones (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003eThe results from the models predicting ERPs amplitude from Category, non-anhedonic BDI-II scores and their interaction are reported in the Supplementary Material (see Supplementary, Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e online). No significant effect of non-anhedonic BDI-II or Category \u003cem\u003e\u0026times;\u003c/em\u003e non-anhedonic BDI-II emerged.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eThe present study examined whether anhedonia influenced the emotional processing of pleasant stimuli at different stages, as assessed with different ERPs during the emotional S1-S2 task. While anhedonia is traditionally described as the inability to feel pleasure, recent perspectives suggest it may be also linked to impairments in motivation and anticipation, rather than solely a lack of pleasure [\u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e7\u003c/span\u003e, \u003cspan citationid=\"CR81\" class=\"CitationRef\"\u003e81\u003c/span\u003e]. In this context and given that emotional processing involves multiple stages (i.e., anticipation, elaboration), understanding how each stage relates to anhedonia may be relevant for the early identification of specific processes involved in depression vulnerability. Notably, in the current sample, higher levels of anhedonia, but not non-anhedonic depressive symptoms, were associated with larger anticipation (SPN) and blunted elaboration (LPP) of pleasant stimuli.\u003c/p\u003e\u003cp\u003eThe results from the LPP are in line with the formulated hypothesis and suggest a reduced elaboration and motivated attention to pleasant stimuli in individuals with higher levels of anhedonia. This finding is consistent with prior research, which has reported blunted LPP in individuals with both clinical [\u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e48\u003c/span\u003e] and subclinical depression [\u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e68\u003c/span\u003e], among those with a familial risk for the disorder [\u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e50\u003c/span\u003e, \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e82\u003c/span\u003e], and as a predictor of future depression onset [83]. Notably, in this study, a reduced LPP to pleasant stimuli was specifically associated with anhedonia rather than non-anhedonic depressive symptoms, suggesting that this component may uniquely reflect anhedonia rather than general depressive symptomatology, at least in individuals without a formal diagnosis of depression.\u003c/p\u003e\u003cp\u003eContrary to the initial hypothesis, the results from the SPN suggest that anhedonia may increase the anticipation of pleasant stimuli. This heightened anticipation could serve as a compensatory mechanism to counterbalance the already diminished elaboration and processing of these stimuli. Instead, in clinical depression both anticipation and elaboration may become impaired, reflecting a more extended deficit of appetitive stimuli processing [\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e, \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e]. Notably, a similar pattern of heightened anticipation has been observed in individuals remitted from a major depressive disorder [\u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e84\u003c/span\u003e] and in those exhibiting depressive symptoms [59] during reward processing tasks. Thus far, this is the only study highlighting this effect during an emotional picture processing paradigm, and further research employing similar tasks may aid in clarifying this pattern as a marker of depression vulnerability as well as its relationship with anhedonia. However, caution is warranted in interpreting this result, as the SPN, typically identified as a slow negative waveform, consistently maintained its amplitude above the isoelectric line across all emotional categories, possibly because the employed paradigm did not activate the appropriate anticipation mechanisms. Furthermore, the valence of the emotional stimuli was always predicted by corresponding cues, and this task might not have sufficiently engaged anticipatory mechanisms during the processing of emotional stimuli. The choice of maintaining congruency between S1 and S2 was made to ensure coherence with previous studies [\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e37\u003c/span\u003e, \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e, \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e43\u003c/span\u003e] but future research should consider incorporating such manipulations to better understand the dynamics of anticipation and emotional processing.\u003c/p\u003e\u003cp\u003eIn addition, neither anhedonia nor non-anhedonic depressive symptoms significantly predicted changes in Cue-P300 amplitude, indicating that these conditions did not influence the initial affective engagement and preparation to elaborate the emotional stimuli. This finding is in contrast with other studies reporting a reduced Cue-P300 in individuals with anhedonia [\u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e54\u003c/span\u003e] or depression vulnerability [\u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e52\u003c/span\u003e]. The discrepancy may stem from differences in experimental paradigms, as prior studies assessing the Cue-P300 often employed reward tasks that typically required a motor response.\u003c/p\u003e\u003cp\u003eTaken together, the larger SPN and smaller LPP to pleasant images suggest a complex pattern of affective processing in individuals with anhedonia, marked by a rapid transition from increased anticipation to reduced elaboration specifically of pleasant stimuli. Although some studies have reported diminished affective responses to both unpleasant and pleasant stimuli in individuals vulnerable to depression [\u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e50\u003c/span\u003e, \u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e82\u003c/span\u003e], indicating a general reduction in motivational drive [\u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e85\u003c/span\u003e], the present findings suggest that anhedonic symptoms may specifically impair the emotional processing of pleasant stimuli.\u003c/p\u003e\u003cp\u003eBefore testing the main hypothesis, the effect of the emotional category was examined. This effect was found to be significant across all ERPs. Specifically, the LPP was larger for both pleasant and unpleasant images compared to neutral ones, indicating heightened elaboration of emotional stimuli. In contrast, the Cue-P300 and SPN were larger for pleasant (but not unpleasant) images relative to neutral ones, reflecting increased affective engagement and anticipation of pleasant stimuli. Most studies have reported increased amplitude of these components during the anticipation of appetitive stimuli using reward tasks [\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e, \u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e53\u003c/span\u003e, \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e55\u003c/span\u003e, \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e57\u003c/span\u003e]. However, the current findings may extend these previous results, suggesting a general tendency to engage with and anticipate appetitive content, even when the pleasant stimuli are passively presented.\u003c/p\u003e\u003cp\u003eFinally, subjective SAM ratings confirmed that emotional pictures were perceived as more arousing than neutral ones, validating the experimental manipulation. Nonetheless, anhedonia did not influence the self-report ratings of valence and arousal. These findings align with other studies suggesting that event-related potentials may be more sensitive than self-report measures in detecting anhedonic symptoms, capturing attentional and emotional processes that are not reflected by self-report ratings [\u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e49\u003c/span\u003e, \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e69\u003c/span\u003e]. This underscores the importance of combining psychophysiological and self-report measures to gain deeper insights into the affective processing patterns underlying vulnerability to depression.\u003c/p\u003e\u003cp\u003eOverall, this study contributes at enhancing the understanding of anhedonia as a risk factor for depression. From the RDoC perspective, both clinical depression and its associated risk have been linked to the hypoactivation of the Positive Valence System (PVS), characterized by reduced anticipation and diminished elaboration of appetitive stimuli [\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e20\u003c/span\u003e, \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e48\u003c/span\u003e, \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e49\u003c/span\u003e, \u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e86\u003c/span\u003e]. However, the present findings indicate that vulnerability to depression associated with anhedonia may involve a more complex pattern of affective processing, marked by increased anticipation followed by reduced elaboration of pleasant stimuli. Further research may aid in examining this pattern in greater detail, to better delineate anhedonic symptoms within the PVS and across other dimensions of the RDoC framework.\u003c/p\u003e\u003cp\u003eFrom a clinical perspective, these findings provide preliminary insights into the mechanisms underlying anhedonia and may help inform potential approaches to targeted screening and interventions. From these findings, it can be suggested that increasing exposure to rewarding experiences or employing behavioral activation techniques could be particularly beneficial for individuals with anhedonia [\u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e87\u003c/span\u003e]. In this context, several clinical interventions have been developed to enhance the activation of the PVS, including Positive Affect Treatment [88], Amplification of Positivity [89], and Behavioral Activation for Anhedonia [\u003cspan citationid=\"CR88\" class=\"CitationRef\"\u003e90\u003c/span\u003e]. These interventions primarily emphasize the active pursuit of pleasant experiences [\u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e91\u003c/span\u003e, \u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e92\u003c/span\u003e] and incorporate exposure to pleasant stimuli, which foster their enhanced elaboration [88,89,90,91,92,93]. Therefore, integrating psychophysiological measures such as ERPs can provide valuable insights into the effectiveness of these interventions. For example, savoring, an emotion regulation technique designed to enhance, prolong, and intensify positive emotions, has been found to significantly increase LPP amplitude during the processing of pleasant stimuli [94,95]. Leveraging ERPs in future research could refine the assessment of depression vulnerability and enhance the specificity of both preventive and clinical interventions.\u003c/p\u003e\u003cp\u003eThese findings must be considered in light of several limitations. First, the sample was confined to young adults, primarily university students who were not completely free from depressive symptoms [\u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e65\u003c/span\u003e]. Second, it was not possible to separate the effects of anhedonia from that of other confounding variables that can impact affective processing, such as anxiety symptoms [\u003cspan citationid=\"CR96\" class=\"CitationRef\"\u003e96\u003c/span\u003e] having had a depressive episode in the past, or parental history of depression [\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e, \u003cspan citationid=\"CR95\" class=\"CitationRef\"\u003e97\u003c/span\u003e]. In addition, the SHAPS was the sole measure of anhedonia included in this study [\u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e63\u003c/span\u003e]. This scale has good convergent and discriminant validity [98] but only encompasses the construct of consummatory anhedonia [\u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e].\u003c/p\u003e\u003cp\u003eIn sum, this study adds to the current literature exploring how anhedonia, a core symptom and vulnerability factor of depression, impacts neural responses to emotional stimuli at different stages of affective processing. Results indicate that individuals with anhedonia are more likely to experience greater anticipation and reduced elaboration of pleasant stimuli. These findings enhance the understanding of depression vulnerability and may inform the development of targeted preventive interventions aimed at regulating the aberrant processing of pleasant stimuli.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003ch2\u003eCompeting interests\u003c/h2\u003e\u003cp\u003eThe authors declare no competing interests.\u003c/p\u003e\u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eV.M., C.D.A, and S.M.B. conceived and designed the study; V.M. and R.M. conducted the study; V.M. analyzed the data; V.M., C.D.A., and S.M.B visualized and interpreted the results; V.M. wrote the first draft; all authors reviewed the manuscript.\u003c/p\u003e\u003ch2\u003eAcknowledgement\u003c/h2\u003e\u003cp\u003eWe acknowledge financial support under the National Recovery and Resilience Plan (NRRP) funded by the European Union \u0026ndash; NextGenerationEU \u0026ndash; Project Numbers: 20228P4H2K and P20223PTH4, adopted by the Italian Ministry of University and Research (MUR).\u003c/p\u003e\u003ch2\u003eData Availability\u003c/h2\u003e\u003cp\u003eThe datasets and materials analyzed in this study are not publicly available due to ethical considerations. 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Psychometric evaluation of the Snaith-Hamilton pleasure scale in adult outpatients with major depressive disorder. \u003cem\u003eInternational clinical psychopharmacology\u003c/em\u003e,\u003cstrong\u003e 25\u003c/strong\u003e(6), 328\u0026ndash;333; https://doi.org/10.1097/YIC. 0b013e32833eb5ee (2010).\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Anhedonia, depression vulnerability, ERPs, S1-S2 task, SPN, LPP","lastPublishedDoi":"10.21203/rs.3.rs-7073207/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7073207/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eAnhedonia, defined as a diminished capacity to experience pleasure form appetitive stimuli, is a core symptom of depression and a predisposing factor for its development. Prior research links anhedonia with blunted emotional processing of appetitive stimuli. Yet, emotional processing encompasses multiple stages (cue engagement, affective anticipation, and elaboration), and how each stage relates to anhedonia remains unclear. This study examined these associations in a sample of university students (\u003cem\u003en\u003c/em\u003e\u0026thinsp;=\u0026thinsp;45, 31 females) with varying levels of anhedonia. Participants underwent electroencephalography recording during an S1-S2 paradigm, in which a cue (S1) anticipated the valence (pleasant, neutral, unpleasant) of an upcoming emotional image (S2). Three event-related potentials (ERPs) were assessed: the Cue-P300 (reflecting cue evaluation and engagement), the Stimulus Preceding Negativity (SPN; reflecting affective anticipation), and the Late Positive Potential (LPP; reflecting affective elaboration). While the LPP was larger for emotional vs. neutral images, the Cue-P300 and the SPN were more pronounced for pleasant (but not unpleasant) vs. neutral stimuli. Notably, anhedonia, independent of other depressive symptoms, was associated with an increased SPN and a blunted LPP for pleasant stimuli. These findings suggest a complex pattern of emotional processing in anhedonia, marked by increased anticipation but reduced elaboration of appetitive stimuli.\u003c/p\u003e","manuscriptTitle":"The time course of affective processing in anhedonia: insights from event-related potentials","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-09-03 05:09:57","doi":"10.21203/rs.3.rs-7073207/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"ee7c1381-c37f-41f8-8479-fa75e80e0276","owner":[],"postedDate":"September 3rd, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[{"id":54022378,"name":"Health sciences/Diseases"},{"id":54022379,"name":"Biological sciences/Neuroscience"},{"id":54022380,"name":"Biological sciences/Psychology"},{"id":54022381,"name":"Social science/Psychology"}],"tags":[],"updatedAt":"2025-10-03T07:09:16+00:00","versionOfRecord":[],"versionCreatedAt":"2025-09-03 05:09:57","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-7073207","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7073207","identity":"rs-7073207","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}