Arousal modulates valence effects on both early and late stages of affective picture processing in a passive viewing task.

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Arousal modulates valence effects on both early and late stages of affective picture processing in a passive viewing task ab

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Chunliang Feng , Wanqing Li , Tengxiang Tian , Yi Luo , Ruolei Gu , Chenglin Zhou & Yued

jia Luo a

School of Kinesiology, Shanghai University of Sport, Shanghai, China

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State Key Laboratory of Cognitive Neuroscience and Learning, Beijing Normal University, Beijing, China c

Key Laboratory of Behavioral Science, Institute of Psychology, Chinese Academy of Sciences, Beijing, China d

Institute of Affective and Social Neuroscience, Shenzhen University, Shenzhen, China Published online: 07 Mar 2014.

To cite this article: Chunliang Feng, Wanqing Li, Tengxiang Tian, Yi Luo, Ruolei Gu, Chenglin Zhou & Yue-jia Luo (2014) Arousal modulates valence effects on both early and late stages of affective picture processing in a passive viewing task, Social Neuroscience, 9:4, 364-377, DOI: 10.1080/17470919.2014.896827 To link to this article: http://dx.doi.org/10.1080/17470919.2014.896827

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SOCIAL NEUROSCIENCE, 2014 Vol. 9, No. 4, 364–377, http://dx.doi.org/10.1080/17470919.2014.896827

Downloaded by [University of Newcastle (Australia)] at 22:09 01 September 2014

Arousal modulates valence effects on both early and late stages of affective picture processing in a passive viewing task Chunliang Feng1,2, Wanqing Li2, Tengxiang Tian2, Yi Luo2, Ruolei Gu3, Chenglin Zhou1, and Yue-jia Luo4 1

School of Kinesiology, Shanghai University of Sport, Shanghai, China State Key Laboratory of Cognitive Neuroscience and Learning, Beijing Normal University, Beijing, China 3 Key Laboratory of Behavioral Science, Institute of Psychology, Chinese Academy of Sciences, Beijing, China 4 Institute of Affective and Social Neuroscience, Shenzhen University, Shenzhen, China 2

Valence and arousal are primary dimensions of affective stimuli. An interaction of these two factors on affective processing is largely unknown. In this study, the processing of affective pictures was investigated in an orthogonal valence (positive vs. negative) by arousal (high vs. low) task design. Participants were instructed to passively view each presented picture and did not need to make any responses. The valence by arousal interaction was observed on three event-related potential (ERP) components, including the P2 (160–190 ms), N2 (220–320 ms) and late positive potential (LPP) (400–700 ms). This interaction revealed that negative pictures evoked larger neural responses compared with positive pictures (i.e., negative bias) at the high-arousal level, whereas negative pictures evoked smaller neural responses than positive pictures (i.e., positive offset) at the low-arousal level. The current results suggest that the effect of emotional valence on affective picture perception is modulated by levels of arousal at both early and late stages of processing. Finally, the main effect of valence was evident in the P1 component (90–110 ms) and arousal effect in the N1 component (120–150 ms).

Keywords: Valence; Arousal; Event-related potential (ERP); Emotion; Positive offset; Negative bias.

Valence and arousal have been widely considered as two primary dimensions of affective stimuli (Barrett & Russell, 1999; Lang, Bradley, & Cuthbert, 1990; Olofsson, Nordin, Sequeira, & Polich, 2008). Valence ranges from pleasure and approach to displeasure and withdrawal, whereas arousal reflects the general activation level evoked by either positive or negative stimuli. The modulations of valence and arousal on affective picture perception have been extensively

investigated by using the event-related potential (ERP) technique in recent decades. A recent comprehensive review posited that valence primarily modulates the early stage (i.e., 200 ms) of processing (Olofsson et al., 2008). Regarding the valence effects, it has been frequently reported that negative pictures elicit larger

Correspondence should be addressed to: Chenglin Zhou, School of Kinesiology, Shanghai University of Sport, Shanghai 200438, China. E-mail: [email protected] The authors thank Dr. Stefan Wiens and Dr. Steve Hillyard for providing helpful comments on the manuscript. This research was supported by the National Basic Research Program of China (973 Program) [grant nos. 2014CB744600 and 2011CB711000] and the National Natural Science Foundation of China [grant nos. 91132704, 31171004, 31300847].

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MODULATIONS OF AROUSAL ON VALENCE EFFECT

early ERP responses (e.g., P1, N1, and P2) compared with positive and neutral pictures, indicating a “negativity bias” at early stages (Carretié, Albert, LópezMartín, & Tapia, 2009; Carretié, Martín-Loeches, Hinojosa, & Mercado, 2001; Smith, Cacioppo, Larsen, & Chartrand, 2003). This negativity bias has been interpreted as a consequence of the rapid allocation of attentional resources to negative stimuli. These urgent responses to negative stimuli are adaptive for individuals and species, such that threatening situations can be detected and avoided quickly (Becker & Byrne, 1985; Olofsson et al., 2008). According to this viewpoint, attention is automatically oriented to threatening stimuli (Öhman & Mineka, 2001). However, several recent studies have observed that positive pictures can evoke comparable or even more pronounced early ERP responses compared with negative pictures (Brown, van Steenbergen, Band, de Rover, & Nieuwenhuis, 2012; Feng, Wang, Wang, Gu, & Luo, 2012; Foti, Hajcak, & Dien, 2009; Lithari et al., 2010). In brief, the current literature has not yet reached a consensus about the direction of valence effects at early stages of stimulus processing (Olofsson et al., 2008). In addition, several studies have also observed valence effects at late temporal stages, such that negative pictures evoke a larger late positive potential (LPP) compared with positive and neutral pictures (Cano, Class, & Polich, 2009; Conroy & Polich, 2007; Delplanque, Silvert, Hot, Rigoulot, & Sequeira, 2006; Ito, Larsen, Smith, & Cacioppo, 1998). These results suggest that negative pictures activate enhanced elaborative processing compared with positive and neutral pictures. However, these late valence effects are not prominent in the current literature. In contrast, numerous studies have observed that late ERP responses (e.g., LPP) are sensitive to the factor of arousal rather than valence (De Cesarei & Codispoti, 2011; Moratti, Saugar, & Strange, 2011; Sabatinelli et al., 2007). It is thus unclear regarding on whether arousal, valence, or both factors together modulate the late ERP components. In our opinion, the discrepancy about valence effects observed in previous ERP studies might be attributed to the modulatory influence of arousal on valence effects (Feng, Wang, Wang, et al., 2012; Gianotti et al., 2008; Lewis, Critchley, Rotshtein, & Dolan, 2007; Robinson, Storbeck, Meier, & Kirkeby, 2004). This viewpoint is consistent with the model of Cacioppo and Berntson (1994). According to this model, the appetitive system tends to respond more strongly compared with the aversive system when the arousal level of affective stimuli is low (positive offset), whereas the reverse is true when the arousal level is high (negativity bias)

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(Cacioppo & Berntson, 1994; Cacioppo, Gardner, & Berntson, 1997; Cacioppo, Gardner, & Berntson, 1999; Ito & Cacioppo, 2001; Ito, Cacioppo, & Lang, 1998; Smith et al., 2003). Therefore, negative stimuli could evoke either higher or lower neural and behavioral responses compared with positive stimuli, depending on the arousal levels of affective stimuli. Recent behavioral and functional magnetic resonance imaging (fMRI) studies have provided support for the model of Cacioppo et al. These studies have consistently observed that arousal modulates valence effects on affective picture processing at both behavioral and neural levels (Eder & Rothermund, 2010; Lewis et al., 2007; Mickley Steinmetz, Addis, & Kensinger, 2010; Nielen et al., 2009; Robinson et al., 2004). Compared with behavioral and fMRI studies, the ERP technique provides fine-grain temporal resolution of mental processing (e.g., Liotti, Woldorff, Perez, & Mayberg, 2000). This feature of ERP recordings may help to reveal the dynamic modulations of arousal on valence effects at distinct temporal stages. However, previous ERP studies employing an orthogonal design have revealed inconsistent findings regarding the valence by arousal interaction. Specifically, many studies did not observe the valence by arousal interaction on either early (e.g., P2) or late ERP (e.g., LPP) components, but instead observed independent effects of valence and arousal (Briggs & Martin, 2009; Lithari et al., 2010; Rozenkrants & Polich, 2008; van Lankveld & Smulders, 2008). In contrast, some other studies have reported the valence by arousal interactions by using the ERP technique (Feng, Wang, Liu, et al., 2012; Gianotti et al., 2008), which are consistent with recent evidence from fMRI and behavioral studies. For instance, a robust valence by arousal interaction has been observed across early and late temporal stages (i.e., P2, N2, and P3) in our recent study (Feng, Wang, Liu, et al., 2012). In that study, we investigated the implicit processing of affective pictures and employed an indirect task that differed from those used in many previous ERP studies, including the studies using an orthogonal design. To examine the reliability of our previous findings, we applied a commonly used paradigm (i.e., passive viewing task) in the current study. The passive viewing paradigm has been frequently used in both early (see also Olofsson et al. (2008)) and more recent studies (e.g., de Rover et al., 2012; Ferrari, Bradley, Codispoti, Karlsson, & Lang, 2013; Leite et al., 2013; Wheaton et al., 2013). In summary, we suggest that valence effects on affective picture processing might be modulated by arousal. The current study aimed to test this hypothesis and replicate the findings of our recent study by employing a classical passive viewing paradigm. In


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light of our previous study, it was hypothesized that valence by arousal interactions would be present in both early and late ERP responses. Specifically, we expected that interactions would arise as early as the P2 component and extend to the N2 and LPP components. The interaction effects in these ERP components would reveal that negative pictures evoke larger ERP responses compared with positive pictures with a high-arousal level, whereas ERP responses associated with negative pictures would be smaller compared with those associated with positive pictures with a low-arousal level (Feng, Wang, Liu, et al., 2012; Gianotti et al., 2008). In addition, a main effect of valence was also expected for the P2 component (Huang & Luo, 2006, 2007), which has been considered as the most robust index of affective picture processing at early stages of affective processing (Carretié, Martín-Loeches, et al., 2001; Carretié, Mercado, Tapia, & Hinojosa, 2001). Finally, a main effect of arousal was expected for the LPP component (Cuthbert, Schupp, Bradley, Birbaumer, & Lang, 2000; Schupp et al., 2007), which is the most reliable index of affective processing at late stages (Bradley, Hamby, Löw, & Lang, 2007; Hajcak, MacNamara, & Olvet, 2010).

METHODS Participants Twenty-five individuals (10 females) aged 19–27 years (mean age = 21.3 years) participated in the present study. All participants had normal or corrected-to-normal vision and did not have any history of psychiatric or neurological illness. This study and the recruitment of participants were approved by the Beijing Normal University Institutional Review Board (IRB). Written informed consents were collected for all participants. A male participant was excluded from the analysis due to massive head movements during the formal task. Therefore, the final sample consisted of 24 participants in total.

Stimuli Pictures used in our study were selected from the International Affective Picture System (IAPS) based on normative ratings of valence and arousal (Lang, Bradley, & Cuthbert, 2001). These ratings were further validated on a local sample of 300 Chinese people (Liu, Xu, & Zhou, 2009). There were four types of stimuli: high-arousing positive (HP), low-

TABLE 1 The mean (with SD) valence and arousal in each condition Positive

Negative

High arousal Low arousal High arousal Low arousal Valence Arousal

6.61 (.56) 6.34 (.36)

6.15 (.49) 4.87 (.31)

1.88 (.38) 6.07 (.18)

2.82 (.62) 4.58 (.46)

arousing positive (LP), high-arousing negative (HN), and low-arousing negative (LN), with 15 pictures for each type (HP: 2216, 4607, 4658, 4687, 5470, 5621, 5629, 8030, 8034, 8090, 8180, 8190, 8370, 8470, 8490; LP: 1500, 1590, 1670, 2080, 2304, 2340, 2360, 2500, 2515, 5220, 7238, 7285, 7430, 7545, 8320; HN: 3053, 3060, 3071, 3102, 3120, 3130, 3170, 3261, 3350, 3400, 3530, 6313, 6570, 9570, 9921; LN: 2205, 2700, 2722, 2750, 2753, 9001, 9041, 9046, 9265, 9280, 9320, 6010, 9330, 9415, 9830). All pictures were adjusted to a size of 12° × 9°. Pictures were presented on a CRT monitor with an 80-Hz refresh rate using the Psychophysics Toolbox extensions in MATLAB (Brainard, 1997; Pelli, 1997). The normative ratings of valence and arousal ratings of the selected affective pictures have been presented elsewhere (Feng, Wang, Liu, et al., 2012). The ratings of valence and arousal from the local sample are displayed in Table 1. It should be noted here that the focus of interest in the current study was on how arousal modulates valence effects in affective picture processing. In this regard, the four types of affective pictures selected for the current study will effectively satisfy this aim. In accord with the current study, many previous ERP studies employing orthogonal design did not use neutral pictures (Lithari et al., 2010; Rozenkrants, Olofsson, & Polich, 2008; Rozenkrants & Polich, 2008).

Procedure Participants sat in a dimly lit and sound-attenuated chamber facing a CRT monitor 80 cm away from their eyes. Each picture was presented for 1000 ms, followed by a intertrial interval of 1700–2300 ms. Participants were instructed to passively view the sequence of pictures (Bradley et al., 2007; Codispoti, Ferrari, & Bradley, 2007; Ferrari et al., 2013; Keil et al., 2002). The whole experiment consisted of 5 blocks containing 60 pictures each. In each block, each of the 60 pictures was presented once, and the sequence of trials in each block was pseudo-random, with the constraints that pictures belonging to the same type


(e.g., high-arousal negative pictures) were not presented in adjacent trials for more than three times.


EEG recording The electroencephalogram (EEG) was recorded from 64 scalp sites using electrodes mounted in an elastic cap (Brain Product, GmbH, Germany), with an online reference to the left mastoid. The horizontal electroencephalogram (HEOG) was recorded with two electrodes placing laterally to the right and left eyes. The vertical electroencephalogram (VEOG) was recorded with electrodes placed above and below the right eye. All interelectrode impedances were maintained below 10 kΩ. The EEG and EOG were amplified using a .01–100 Hz band-pass and continuously sampled at 500 Hz in each channel for off-line analysis. All EEG signals were re-referenced off-line to the average of left and right mastoids. It is worth noting that averaged mastoid references have been recommended as the standard in the field of cognitive neuroscience, especially when a large array of electrodes is not used (for detailed discussion, see also Luck (2005)). The EEG data were low-pass filtered below 30 Hz (24 dB/oct) and were corrected for eye movements or blinks using the Gratton and Coles method (Gratton & Coles, 1989) implemented in the Brain Vision analysis software (Brain Product, GmbH, Germany). Trials containing EEG sweeps with amplitudes exceeding ±80 μV were excluded.

Data reduction and analysis ERPs elicited by the pictures were averaged over epochs of 1000 ms, with a 200-ms prestimulus baseline. Five different ERP components, including the P1, N1, P2, N2, and LPP, were measured in the current study (Keil et al., 2002; Olofsson & Polich, 2007; Rozenkrants et al., 2008). Different sets of electrodes were chosen for the mean amplitude measurements of these components. Specifically, occipitotemporal (PO3/PO4) and occipital (O1/O2) electrodes were chosen for the P1 component (90–110 ms) and the N1 component (120–150 ms); frontal (F3/Fz/F4), fronto-central (FC3/FCz/FC4), and central (C3/Cz/ C4) electrodes for the P2 component (160–190 ms) and the N2 component (220–320 ms); and frontal (F3/ Fz/F4), central (C3/Cz/C4), and parietal (P3/Pz/P4) electrodes for the LPP component (400–700 ms). The mean amplitudes of these components over the indicated intervals were then analyzed in repeated measures, ANOVAs, with the factors of valence

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(positive vs. negative), arousal (high vs. low), hemisphere (left vs. middle vs. right), as well as electrode as within-subject factors. The effects of hemisphere and electrode were reported only when they interacted with valence or arousal (Briggs & Martin, 2009; van Lankveld & Smulders, 2008; Van Strien, Langeslag, Strekalova, Gootjes, & Franken, 2009). P-values were corrected for deviations using Greenhouse-Geisser correction if necessary. Bonferroni correction was used for multiple comparisons with alpha (α) = .05. It should be noted that the Bonferroni correction was used to correct the number of contrast pairs for each factor (Figure 1).

RESULTS P1 component A four-way repeated measures ANOVA of valence (negative vs. positive) × arousal (high vs. low) × hemisphere (left vs. right) × electrode (occipitotemporal vs. occipital) on P1 amplitude revealed a significant main effect of valence (F(1, 23) = 5.66, p < .05, η2p =.20), such that negative pictures elicited a larger P1 amplitude than positive pictures (Bonferroni post hoc test, p < .05) (Figure 2).

N1 component A four-way repeated measures ANOVA of valence (negative vs. positive) × arousal (high vs. low) × hemisphere (left vs. right) × electrode (occipitotemporal vs. occipital) on N1 amplitude revealed a significant main effect of arousal (F(1, 23) = 8.34, p < .01, η2p =.27), such that high-arousing pictures elicited larger N1 amplitude than low-arousing pictures (Bonferroni post hoc test, p < .05).

P2 component A four-way repeated measures ANOVA of valence (negative vs. positive) × arousal (high vs. low) × hemisphere (left vs. middle vs. right) × electrode (frontal vs. fronto-central vs. central) on P2 amplitude revealed a significant main effect of arousal (F(1, 23) = 5.51, p < .05, η2p =.19), such that high-arousing pictures evoked a larger P2 than low-arousing pictures. The valence × arousal interaction was also significant (F(1, 23) = 30.10, p < .0005, η2p =.57). Bonferroni post hoc test revealed that negative pictures elicited a larger P2 than positive pictures at high-arousal level (p < .05), whereas negative


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Figure 1. The grand average ERPs evoked by high-arousing positive, low-arousing positive, high-arousing negative, and low-arousing negative pictures. Occipitotemporal and occipital electrodes (green) were chosen for P1 and N1 measurement; frontal, fronto-central, and central electrodes (included in the blue ellipse) were chosen for P2 and N2 measurement; frontal, central, and parietal electrodes (red) were chosen for LPP measurement.

Figure 2. The amplitudes of P1 and N1 (error bars show 1 SE). P1 amplitudes in positive and negative conditions (i.e., data from low- and high-arousing conditions were collapsed) are illustrated; N1 amplitudes in low- and high-arousing conditions (i.e., data from positive and negative conditions were collapsed) are illustrated.


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Figure 3. The amplitudes of P2, N2, and LPP evoked by high-arousing positive, low-arousing positive, high-arousing negative, and lowarousing negative pictures (error bars show 1 SE).

pictures elicited a smaller P2 than positive pictures at low-arousal level (p < .05) (Figure 3).

N2 component A four-way repeated measures ANOVA of valence (negative vs. positive) × arousal (high vs. low) × hemisphere (left vs. middle vs. right) × electrode (frontal vs. fronto-central vs. central) on N2 amplitude yielded significant main effects of valence (F(1, 23) = 6.42, p < .05, η2p =.22) and arousal (F(1, 23) = 16.24, p < .005, η2p =.41). These main effects revealed that negative pictures evoked a larger positivity in the N2 window than positive pictures (p < .05), while higharousing pictures elicited a larger positivity in the N2 window than low-arousing pictures (p < .05). In addition, the valence × arousal interaction was significant (F(1, 23) = 38.40, p < .0005, η2p =.63). Bonferroni post hoc testing revealed that negative pictures elicited a larger positivity in the N2 window than positive pictures at high-arousal level (p < .05), whereas negative pictures elicited a smaller positivity in the N2 window compared with positive pictures at low-arousal level (p < .05).

LPP component A four-way repeated measures ANOVA of valence (negative vs. positive) × arousal (high vs. low) × hemisphere (left vs. middle vs. right) × electrode (frontal vs. central vs. parietal) on LPP amplitude revealed that negative pictures elicited a larger LPP amplitude than positive pictures, as indicated by a significant main effect of valence (F(1, 23) = 62.05, p < .05, η2p =.73). High-arousing pictures elicited a larger LPP amplitude than low-arousing pictures, as shown by a significant main effect of arousal (F(1, 23) = 111.93, p < .0005, η2p =.83). Most importantly, the valence × arousal interaction was significant (F(1, 23) = 70.53, p < .0005, η2p =.75), such that negative

pictures elicited a larger LPP amplitude than positive pictures at high-arousal level (p < .05), whereas negative pictures elicited a smaller LPP amplitude than positive pictures at low arousal (p < .05). The valence × arousal × hemisphere × electrode interaction was also significant (F(4, 92) = 3.72, p < .05, η2p =.14). Bonferroni post hoc testing revealed that high-arousing negative pictures elicited a larger LPP amplitude than high-arousing positive pictures over both hemispheres (p < .05), whereas low-arousing negative pictures elicited a smaller LPP amplitude than low-arousing positive pictures over frontal and central electrodes (p < .05), but not over parietal electrodes (p > .05) (Figure 4).

DISCUSSION The current study investigated how arousal modulates valence effects on affective picture processing and the time course of these modulations. The main effect of valence was observed in the P1 component (90–110 ms) and the main effect of arousal in the N1 component (120–150 ms). The valence by arousal interaction was observed across early and late temporal stages, including P2 (160– 190 ms), N2 (220–320 ms), and LPP (400– 700 ms). Noteworthy, the valence by arousal interaction on ERP components revealed that negative pictures evoked larger neural responses than positive pictures in the high-arousing condition, whereas the reverse was true for the low-arousing condition. In this regard, it is unlikely that the interaction pattern on ERP components could be attributed to the larger differences in valence ratings between positive and negative pictures for high-arousing condition than low-arousing condition (i.e., 4.73 vs. 3.33, see Table 1). In the following sections, we discuss the roles of valence and arousal in affective picture processing at each temporal stage.


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Figure 4. Scalp topography of ERPs evoked by high-arousing positive (HP), low-arousing positive (LP), high-arousing negative (HN), and low-arousing negative (LN) pictures on P1 (90–110 ms), N1 (120–150 ms), P2 (160–190 ms), N2 (220–320 ms), and LPP (400–700 ms). Please note that the topographies referred to mastoid references and that the scales were different between ERP components.

The early temporal stages (before 200 ms) Previous studies have reported that negative stimuli evoke an enhanced P1 component compared with positive and neutral stimuli, suggesting a negativity bias on this component (Delplanque, Lavoie, Hot, Silvert, & Sequeira, 2004; Sass et al., 2010; Smith et al., 2003; Taake, Jaspers-Fayer, & Liotti, 2009). However, other studies have failed to replicate this effect but observed instead that positive and negative pictures elicit comparable P1 amplitudes (Brown et al., 2012; Olofsson & Polich, 2007; Pérez-Edgar & Fox, 2003). In the current study, negative pictures elicited a larger P1 compared with positive pictures. These results support previous studies showing that negativity bias could be present on the P1 component. In contrast, an influence of arousal rather than valence was observed on the N1 component in the current study. As with the other early ERP components, previous studies also report heterogeneous results regarding the N1 component. On the one

hand, some studies have revealed that the N1 is sensitive to the emotional valence of affective pictures (Carretié, Hinojosa, & Mercado, 2003). On the other hand, other studies have found that negative and positive pictures evoke comparable N1 amplitudes (Foti et al., 2009; Hot Saito, Mandai, Kobayashi, & Sequeira, 2006; Schupp, Junghöfer, Weike, & Hamm, 2003; Weinberg & Hajcak, 2010). The current results for the N1 component provide support to the arousal effects observed in previous studies. These findings suggest that valence effects may not be predominant in the early stage of affective picture processing. As with the P1 and N1 components, P2 is also considered to reflect a rapid and coarse attentional resource allocation at early stages of affective picture processing (Carretié, Martín-Loeches, et al., 2001; Carretié, Mercado, et al., 2001; Huang & Luo, 2007). The P2 is a reliable index of negativity bias, such that negative pictures frequently elicit a larger P2 compared with both positive and neutral pictures (Carretié, Ruiz-Padial, López-Martín, & Albert,



2011; Delplanque et al., 2004; Huang & Luo, 2007; Olofsson & Polich, 2007). However, some studies have observed that a positive stimuli evoke a larger P2 component compared with negative stimuli, suggesting that the positive offset can be reflected on the P2 as well (Lithari et al., 2010; Spreckelmeyer, Kutas, Urbach, Altenmüller, & Münte, 2006). Furthermore, several studies have observed an anxiety by valence interaction on the P2 component, such that negative pictures evoked a larger P2 compared with positive pictures in high-anxiety individuals; whereas the reverse was true in low-anxiety individuals (Mercado, Carretié, Hinojosa, & Penacoba, 2009; Mercado, Carretié, Tapia, & Gómez-Jarabo, 2006). These results suggest that high-anxiety individuals may respond more intensively to affective stimuli (i.e., higher level of arousal) compared with low-anxiety individuals (Becker, Rinck, Margraf, & Roth 2001; Sass et al., 2010). In sum, both negativity bias and positive offset have been previously observed on the P2 component, and an individual’s arousal level in response to affective stimuli may determine which effect is predominant. In accord with this account, the current study observed a valence by arousal interaction on the P2, such that negative pictures elicited a larger P2 compared with positive pictures (negativity bias) at high-arousal level, whereas negative pictures elicited a smaller P2 compared with positive pictures (positive offset) at low-arousal level. The current results thus provide direct support to the assertion that arousal ratings of affective pictures determine whether negativity bias or positive offset will be observed on the P2. In short, the current study observed flexible modulations of valence and arousal at early processing stages (i.e., before 200 ms), in which modulations of valence and arousal can be observed even if attentional resources were deviated from affective stimuli (see also Carretié, Hinojosa, Martín‐Loeches, Mercado, & Tapia (2004)). These effects were reflected by three ERP components: the P1, N1, and P2. Specifically, valence effect was initially present on the P1; followed by an arousal effect on the N1; finally there was a valence by arousal interaction on the P2. These results indicate that not only valence but also arousal and the valence by arousal interaction modulate affective picture processing at early temporal stages (Feng, Wang, Liu, et al., 2012). These findings were in accord with the viewpoint that valence effects at early stages are heterogeneous (for a review, see Olofsson et al. (2008)). Importantly, these results are also consistent with some recently proposed models of affective picture processing, which commonly postulate that affective pictures are

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processed in a “coarse-to-fine” way, such that valence information is processed firstly, then arousal information, and finally the comprehensive information (e.g., the valence by arousal interaction in the current study) (Luo, Feng, He, Wang, & Luo, 2010; Weinberg & Hajcak, 2010). Notably, all these effects were detected at early stages in the current study. The current results thus suggest that the neural discrimination of emotional information is much faster than these models have assumed. In addition, the valence by arousal interaction extended to middle (N2, 220–320 ms) and late (LPP, 400–700 ms) temporal stages, as discussed in the following sections.

The middle temporal stage (220–320 ms) Several studies have demonstrated that the neural responses in the middle temporal stage (i.e., N2) primarily reflect automatic processing of affective pictures (Codispoti et al., 2007; Olofsson et al., 2008), whereas other studies have indicated that both automatic and controlled processes modulate the neural responses in this stage (Carretié et al., 2004; Daffner et al., 2000). In contrast to early ERP components, previous studies have consistently revealed arousal effects on the N2, presumably reflecting the selective attention allocated to affective stimuli with intrinsic relevance (i.e., high-arousing pictures) (Olofsson et al., 2008). The current results were consistent with previously reported arousal effects in that high-arousing pictures elicited a larger positivity in the N2 window compared with low-arousing pictures (Amrhein, Mühlberger, Pauli, & Wiedemann, 2004; Cuthbert et al., 2000; Olofsson & Polich, 2007; Rozenkrants & Polich, 2008). The increased positivity in the N2 window is often interpreted as enhanced processing within this window (Amrhein et al., 2004; Rozenkrants & Polich, 2008), and a similar effect and interpretation has also been evidenced in early posterior negativity (EPN, which shared similar latency with N2 component) at centro-medial electrodes (Codispoti et al., 2007; Schupp, Junghöfer, Weike, & Hamm, 2004; Schupp, Markus, Weike, & Hamm, 2003). More importantly, a robust valence by arousal interaction was also evident on the N2. Specifically, negative pictures elicited a larger positivity in the N2 window compared with positive pictures at high-arousal level; whereas negative pictures elicited a smaller positivity in the N2 window compared with positive pictures at low-arousal level. These effects on the N2 were unexpected, since most previous studies have found out that the arousal effect is predominant on


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the N2. However, it is worth noting that previous studies have shown heterogeneous findings about valence effects on the N2. For instance, several studies have found that negative pictures evoke a larger positivity in the N2 window compared with positive pictures (Carretié et al., 2004; Pérez-Edgar & Fox, 2003; Schupp et al., 2004), whereas Lithari et al. (2010) observed that the positivity in the N2 window was smaller for negative pictures compared with positive pictures (Lithari et al., 2010). These studies suggest that the valence effects can also modulate the N2, which is consistent with the current results. In addition, the valence by arousal interaction observed on the N2 in the present study provides a possible resolution of the contradictory results from previous studies.

The late temporal stages (400–700 ms) The LPP component has been considered as the most robust and reliable index of resource allocation at late stages of affective processing (Cuthbert et al., 2000; Hajcak et al., 2010; Hajcak & Olvet, 2008). Presumably, the LPP reflects the sustained modulations of subcortical motivational systems (e.g., the amygdala) on the visual system (Lang, Bradley, & Cuthbert, 1997; Sabatinelli et al., 2007; Sabatinelli, Bradley, Fitzsimmons, & Lang, 2005; Sabatinelli, Lang, Bradley, Costa, & Keil, 2009). Additionally, LPP responses are also subject to top-down modulations from the prefrontal cortex (Carretié, Hinojosa, Albert, & Mercado, 2006; Hajcak et al., 2010; Moratti et al., 2011). In sum, the LPP reflects the combination of automatic and controlled processing of affective pictures (Ferrari, Codispoti, Cardinale, & Bradley, 2008; Moratti et al., 2011). Numerous studies have observed that positive and negative pictures evoke comparable LPP amplitudes, indicating that it is relatively insensitive to emotional valence (Codispoti et al., 2007; De Cesarei & Codispoti, 2011; Moratti et al., 2011; Sabatinelli et al., 2007). The relationship between the LPP and arousal has been supported by evidence that LPP amplitudes correlated with the arousal ratings of affective pictures (Cuthbert et al., 2000). In some studies, however, valence effects were observed on the LPP component even after controlling the arousal ratings of affective pictures (Cano et al., 2009; Conroy & Polich, 2007; Delplanque et al., 2006; Ito, Larsen, et al., 1998). In the current study, main effects of both arousal and valence were observed. These findings are consistent with previous arousal and valence effects on LPP, respectively. More importantly, a valence by arousal interaction was also observed on the LPP,

which showed a similar pattern with the P2 and N2. These novel findings extended previous ERP studies within the framework of Cacioppo and Berntson (1994) in that not only negativity bias but also positive offset could be observed on the LPP. In accord with the current findings, previous fMRI studies have also observed a valence by arousal interaction in both the prefrontal cortex and subcortical motivational systems (Lewis et al., 2007; Mickley Steinmetz et al., 2010; Nielen et al., 2009). For instance, the effective connectivity between the amygdala and inferior frontal gyrus as well as middle occipital gyrus was larger for high-arousing negative and low-arousing positive pictures than low-arousing negative and high-arousing positive pictures. Such neural patterns were associated with recognition performance after fMRI scanning (Mickley Steinmetz et al., 2010). These findings indicate that arousal modulates valence effects on the memory-related processing (i.e., encoding) of affective stimuli (Gomes, Brainerd, & Stein, 2013). Given that LPP responses have also been associated with memory encoding processes (Carretié et al., 2011; Dolcos & Cabeza, 2002; Olofsson et al., 2008) and are linked to brain regions in which valence by arousal interaction has been observed (Liu et al., 2012; Sabatinelli, Lang, Keil, & Bradley, 2007), it is reasonable to conclude that arousal modulates valence effects on LPP. Finally, a comparison between current study and a recent study is found in Feng, Wang, Liu, et al. (2012). The current study generally replicated our previous findings on valence by arousal interaction across early and late temporal stages. In particular, the interaction was observed on P2 and N2 in both studies. Our recent study also observed the interaction on the P3 over parietal electrodes, but the LPP was extensively attenuated by the irrelevant task used in that study, and no differences between conditions were apparent (Feng, Wang, Liu, et al., 2012). In contrast, the interaction was evident on the LPP in the current study with a passive viewing paradigm. Noteworthy, the valence by arousal interaction on the LPP was more pronounced over frontal and central electrodes than the parietal electrodes in the current study. In other words, although our previous study observed valence by arousal interaction across frontal (P2, N2) and parietal (P3) electrodes, the interaction was limited to frontal and central (P2, N2, and LPP) electrodes in the current study. The reason for this discrepancy between the two studies is unclear. In addition, the valence effect in the P1 and the arousal effect in the N1 were not observed in our previous studies. Although affective modulations have been observed on these early components (Carretié et al.,


2004) when participant’s attention is deviated from affective pictures, previous findings are still heterogeneous (Olofsson et al., 2008). Therefore, we suggest that differences in P1 and N1 results between the two studies might partly reflect effects of task. That is, participant’s attention was focused on the affective pictures to a larger extent in the current study than in our previous study.


LIMITATIONS A possible limitation of the current study is that the affective effects observed in the ERPs might be confounded by physical characteristics (e.g., complexity) of pictures. However, this limitation is not specific to the current study but is a common issue for all studies using IAPS pictures, since the vastly diverse physical attributes of these pictures make it difficult to control physical factors when investigating affective effects (see also Flaisch & Schupp (2013)). The influence of physical characteristics on affective modulations has hardly been systematically assessed, and it is difficult to draw a conclusion based on the current literature. For instance, several studies have provided evidence showing that physical features primarily influence affective effects on early ERP components but not on late components (Bradley et al., 2007; but see Cano et al. (2009) and De Cesarei and Codispoti (2006)), whereas others have reported that affective effects at early temporal stages are unaffected by physical attributes (Carretié et al., 2004). To make this issue more complex, it has been recently reported that physical features modulate affective effects at early and late stages in different ways, such that physical attributes suppress early affective effects but enhance late effects (Wiens, Sand, & Olofsson, 2011). The potential confounds of physical attributes in current findings deserve comprehensive investigation in future studies. Another concern is that habituation/repetition effects might contribute to the current findings. Previous studies have consistently revealed that ERP responses to stimuli are modulated by habitation effects at both early (Feng, Luo, & Fu, 2013; Fu Feng, Guo, Luo, & Parasuraman, 2012) and late stages (Codispoti, Ferrari, & Bradley, 2006; Codispoti et al., 2007). In addition, a repetition by emotional content interaction on the N1 component was observed in a study by Carretie et al. (2003), such that negative pictures showed stronger resistance to habitation compared with positive and neutral pictures. However, many recent studies have failed to replicate these findings and instead report that valence

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and arousal effects are unaffected by habituation across both early and late temporal stages (Codispoti et al., 2006, 2007; Olofsson & Polich, 2007; Rozenkrants et al., 2008). Therefore, we suggest it is difficult to determine whether the current findings are influenced by habituation. While the current study focused on the general effects of arousal and valence as did many previous studies, it is worth noting that many other factors such as gender (Syrjänen & Wiens, 2013) and anxiety (Mercado et al., 2009) could modulate affective picture processing. The modulations of these factors were not considered in this study and deserve elaborate investigation.

CONCLUSION The current study investigated how arousal modulates valence effects on affective picture processing in a passive viewing paradigm. To this end, an orthogonal design of valence (positive vs. negative) and arousal (high vs. low) was employed. The current study revealed a main effect of valence on the P1 (90– 110 ms) and an arousal effect on the N1 (120– 150 ms). More importantly, the valence by arousal interaction was observed on the P2 (160–190 ms), N2 (220–320 ms), and LPP (400–700 ms). These results indicate that arousal modulates the valence effects in both coarse and elaborative processing of affective pictures. In conclusion, the modulations of arousal on valence effects are evident across both early and late temporal stages. Original manuscript received 23 April 2013 Revised manuscript accepted 17 February 2014 First published online 7 March 2014

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Incidental self-processing modulates the interaction of emotional valence and arousal.

Modulation of the initial light reflex during affective picture viewing.

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Positive and negative affective processing exhibit dissociable functional hubs during the viewing of affective pictures.

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