BR A I N R ES E A RCH 1 4 18 ( 20 1 1 ) 8 3 –92 Available online at www.sciencedirect.com www.elsevier.com/locate/brainres Research Report N400 and LPP in spontaneous trait inferences Kris Baetens a,⁎, 1 , Laurens Van der Cruyssen a , Anja Achtziger b , Marie Vandekerckhove a , Frank Van Overwalle a,⁎ a Vrije Universiteit Brussel, Belgium Universität Konstanz, Germany b A R T I C LE I N FO AB S T R A C T Article history: Past research on spontaneous trait inferences using event related potentials (ERPs) has Accepted 29 August 2011 consistently reported increased late positive potential (LPP) amplitudes following social Available online 2 September 2011 expectancy violations, but no N400 modulation. In the present study, participants read scenarios describing behaviors of unknown actors. They entailed descriptions of several Keywords: positive trait implying behaviors, followed by a single final sentence describing behavior ERP that was either consistent or inconsistent with the previously implied trait. As in previous N400 studies, we found significantly increased LPP amplitudes following inconsistent behaviors LPP at multiple frontal sites. Unlike in previous research, we also found increased N400 Trait inference amplitudes at several centro-parietal sites. The divergence of these results is explained World knowledge from minor differences in the stimulus presentation procedure and possible overlap of ERP components of opposite polarity. Temporal principal component analysis (PCA) confirmed the separate influence of concurrent LPP and N400 ERP modulations, and the source of the largest factors was located using sLORETA. It is suggested that the increased N400 in response to trait inconsistencies reflects difficulties in understanding unanticipated behavior, while the LPP effect might reflect evaluative incongruence. © 2011 Elsevier B.V. All rights reserved. 1. Introduction Imagine a co-worker you have always known as a particularly honest and ethical person. One day, you haphazardly stumble upon him while he is prying money from the faculty's charity donation box. This probably sets a number of psychological reactions in motion. Initially, you might have difficulties understanding what he is doing because you don't expect somebody to do such a thing, especially not this person. Once you do realize what he is doing, you might be shocked by the discrepancy between his past and present behavior. Finally, you may adjust your expectations and associate him with a negative personality trait like “greedy” or “dishonest”. Indeed, folk wisdom implores us to use observations of past behavior to predict future behavior: “Fool me once, shame on you; fool me twice, shame on me.” Stripped-down, abstracted versions of scenarios such as described above have been used to study event-related potentials (ERPs) associated with trait inference processes (Bartholow et al., 2001, 2003; Van Duynslaeger et al., 2007, 2008; Van Overwalle et al., 2009). In these studies, participants read behavioral descriptions of (unknown) individuals which strongly implied a certain personality trait. Subsequently, one or more critical sentences followed, either consistent or inconsistent with the previously implied trait. All of these studies report a positivity ⁎ Corresponding authors at: Faculty of Psychology and Educational Sciences, Vrije Universiteit Brussel, Pleinlaan 2, 1050 Elsene, Belgium. Fax: +32 2 629 24 89. E-mail addresses: [email protected] (K. Baetens), [email protected] (F. Van Overwalle). 1 PhD fellow of the Research Foundation – Flanders (FWO). 0006-8993/$ – see front matter © 2011 Elsevier B.V. All rights reserved. doi:10.1016/j.brainres.2011.08.067 84 BR A IN RE S E A RCH 1 4 18 ( 20 1 1 ) 8 3 –92 peaking around 500 ms post stimulus in the grand averages of ERPs following the last (critical) word of inconsistent sentences relative to consistent sentences. This peak was interpreted as a late positive potential (LPP) or P300 component (Bartholow et al., 2001), reflecting context-updating processes, and the latency of this potential was considered an index of the process of trait inference (Van Duynslaeger et al., 2007). The trait inconsistencies in the cited ERP studies contrasted with respect to valence and induced affective responses with the previously presented behavior descriptions. Therefore, one might alternatively interpret these late positivities as a consequence of change in valence which triggers increased attention due to the intrinsic motivational significance of stimuli containing emotional information (Holt et al., 2009). Research on emotional word comprehension has established a general association of the processing of emotional information conveyed by words with a late, extended, attention-modulated process encoding their valence, expressed in the LPP component (for a review, see Hacjak et al., 2010). Stimuli of negative valence generally attract more attention than positive stimuli, a tendency termed the “negativity bias” (e.g., Ito et al., 1998). Furthermore, research has shown that evaluative incongruence, a mismatch between the valence of a category and a subsequent target stimulus, is associated with an increased positivity in the ERP that peaks between 300 and 600 ms post stimulus (Cacioppo et al., 1993, 1996). However, is this the only component one should expect after these trait inconsistencies? In what follows, we will propose that modulation of another ERP component, the N400, is also to be expected in paradigms involving disconfirmations of trait inferences. (Bartholow et al., 2001, 2003; Van Duynslaeger et al., 2007, 2008; Van Overwalle et al., 2009). After all, trait inconsistencies could be considered a specific type of world knowledge violation (e.g., you consider this person to be honest, but all of a sudden it turns out he is very sneaky). Moreover, if a person is described as behaving contrary to expectations, surely this information seems more difficult to integrate with the context than if he acts as usual, which should lead to a relative increase in N400 amplitude. Indeed, increased N400 amplitudes have been found in response to violations of stereotypes, another type of person schema (Van Berkum et al., 2008; White et al., 2009). In contrast to these studies, an earlier study by Osterhout et al. (1997) found no increased N400 amplitudes, but instead increased late positivities at about 600 ms (P600) in response to gender stereotype violations (e.g., “The lumberjack sharpened her axe.”). A possible explanation is that the participants interpreted these sentences as containing a grammatical error in applying the pronoun (reliably associated with the P600). The authors went on to predict that “Anomalies involving social categories that are not marked in the grammar (e.g., race) should not elicit the P600 effect but might elicit the N400 effect associated with semantic/pragmatic aspects of language.” (p. 282). Following up on this interpretation, Bartholow et al. (2001, 2003) included a semantically inconsistent condition in order to compare the effects induced by their trait inconsistencies with a classic N400 effect. However, as noted earlier, they found a very dissimilar P300/LPP following trait inconsistencies, and a modulation of the N400 only for semantic inconsistencies. How can the consistent absence of any N400 effect in the trait ERP literature be explained? 1.2. 1.1. The absence of N400 effects The N400 and expectancy violations The N400 component is a negative deflection in the ERP about 400 ms post stimulus. It is a function of semantic inconsistency (e.g., Kutas and Hillyard, 1980) and reflects the ease of integrating a stimulus with a given context, not only for verbal stimuli (Kutas and Federmeier, 2000), but also for other modalities, for example, the comprehension of visually presented real-world events (Sitnikova et al., 2008). The degree of semantic consistency is not a stable quality of any given combination of stimuli, but rather determined flexibly and under influence of the local context (Filik and Leuthold, 2008; Nieuwland and Van Berkum, 2006). For example, “The peanut is in love” will cause a larger N400 than “The peanut is salted”, but the opposite is true if this sentence is preceded by the title “The amorous peanut”. It has been demonstrated that the N400 may also reflect the violation of objective knowledge about the world (Hagoort et al., 2004). Much of the extensive N400 literature can be conveniently organized around the concept of cloze-probability. This is the probability that people will use a given word to complete a sentence, and it is inversely related to the N400 amplitude following the presentation of that word (Federmeier and Laszlo, 2009). Personality traits can be conceptualized as a schema, a type of world knowledge we construct and constantly adapt to anticipate or predict the behaviors of others (Kressel and Uleman, 2010; Read, 1987). Therefore, one might expect to find N400 effects in the aforementioned studies on trait inferences An obvious reason why no previous ERP study on trait inferences has reported modulation of the N400 might be that it was overshadowed by a large positive potential. All previous ERP studies on trait inferences report a large LPP/P300 (Bartholow et al., 2001, 2003; Van Duynslaeger et al., 2007, 2008; Van Overwalle et al., 2009). Positivities occurring simultaneously with the N400 may obscure it (Franklin et al., 2007; Roehm et al., 2007). In such cases, principal component analysis (PCA) may serve to disentangle the influence of simultaneously occurring ERP components (Franklin et al., 2007). The stimuli presentation procedure of previous ERP studies might be an additional factor working against N400 modulation. In all but one of these studies (Bartholow et al., 2001, 2003; Van Duynslaeger et al., 2007, 2008; but see Van Overwalle et al., 2009) expectancy establishing sentences were followed by two critical consistent and two critical inconsistent sentences, presented in a random order. This yielded six possible sequences, as presented in Table 1. As can be seen, only 50% of the inconsistent sentences were unambiguously inconsistent, in that they were not preceded by another inconsistent sentence. Thus, the remaining 50% of the inconsistent sentences were ambiguous (e.g., if John discloses a trusted secret after stealing charity money, is this really inconsistent?). This could have significantly reduced N400 amplitudes for the inconsistent condition, as it constitutes a form of priming (Federmeier and Laszlo, 2009), dramatically changing the local context (Filik and Leuthold, 2008; Nieuwland and Van Berkum, 2006). The 85 BR A I N R ES E A RCH 1 4 18 ( 20 1 1 ) 8 3 –92 Table 1 – Possible sequences of two consistent (C) and inconsistent (I) sentences. Sequence C C C I I I C I I C C I Unambiguous C I C I C I C I I C I C C case is even more problematic for consistent sentences, of which only 25% were unambiguously consistent, that is, not preceded by an inconsistent sentence. Therefore, on average 75% of the consistent sentences were actually ambiguous (e.g., John refusing to disclose the secret after stealing charity money). This may have significantly increased the N400 amplitudes for the consistent condition. Taken together, it is likely that this way of presenting stimuli diminishes possible differences in N400 amplitude between the conditions. Of course, the same reasoning holds for the LPP in the previously mentioned ERP studies on trait inferences. However, assuming that the LPP reflects evaluative incongruence in these paradigms, it is conceivable that this LPP would be less diminished by this procedure. Whereas the expectations about the described actor can be altered drastically by one inconsistent sentence, the valence of the behaviors is not much affected by it (e.g., when you catch John stealing charity money again, this is less surprising but not less upsetting). As such, the evaluative incongruence in the behavior descriptions associated with the LPP remains obvious and therefore more stable. After all, there was always a small minority of one or two sentences that was evaluatively incongruent with the established context and the other critical sentences, making it plausible that they should still stand out. Note that the semantically incongruent sentences in the work of Bartholow et al. (2001, 2003) were similarly randomly shuffled with the trait consistent and inconsistent sentences, and nevertheless followed by a clear increased N400. However, these sentences were semantically incongruent by themselves (e.g., “Gormak gave the stranger a rain”), rather than incongruent with a preceding context. Therefore, one might expect their meaning to be minimally affected by the context in which they were presented. To the best of our knowledge, there has been only one ERP study on trait inferences in which the person description was followed by a single final sentence (Van Overwalle et al., 2009). Again, the authors reported a P300 effect and no N400 effect. However, the statistical analysis employed (negative peaks were only compared in a 50–300 ms window) wasn't ideal for capturing N400 effects, since they should occur later (Kutas and Federmeier, 2000). Interestingly, these authors did report an unexpected significant effect at the Pz site, where the positive peak amplitude was higher for consistent than for inconsistent trait-implying sentences in the 300–450 ms window, which could reflect an N400 modulation. However, since this study had a complex design (analyzing the joint effect of trait and goal inferences) and effects were small, it is difficult to draw clear conclusions from it. Unambiguous I 1st 2nd 1st 2nd Yes Yes Yes No No No 4/12 (25%) Yes No No No No No Yes Yes Yes Yes Yes Yes 6/12 (50%) No No No No No No 1.3. The present research To avoid the methodological limitations of earlier trait inference studies, we presented person descriptions that were each followed by one critical sentence that was either consistent (50% of the trials) or inconsistent (50%) with it. The descriptions implied positive traits, and inconsistencies described behavior triggering a negative trait inference. This choice was based on the finding that negative social behaviors produce stronger inconsistency effects (Bartholow et al., 2001; Van Duynslaeger et al., 2008). In contrast to earlier research on trait inferences using a random shuffling of consistent, inconsistent and irrelevant sentences, we expect that by using single critical sentences, we will obtain an increased N400 following inconsistencies, in line with the prediction of Osterhout et al. (1997). Specifically, we expect that the mean amplitude for inconsistent endings will be more negative than for consistent endings at centro-parietal sites in the 300– 450 ms interval. We additionally expect to find more positive amplitudes following inconsistent endings than following consistent endings in the later 450–1000 ms time interval reflecting evaluative inconsistency, in line with previous trait studies that reported an increased P300/LPP. To disentangle possible simultaneous influences of these components, we will employ principle component analysis (PCA) (Dien et al., 2005). As a first step to shed light on their functional correlates, we will roughly localize the neural sources of important components using sLORETA (Pascual-Marqui, 1999; Pascual-Marqui et al., 1994). 2. Results Fig. 1 displays the grand averages for three selected midline channels (Fz, Cz, Pz). Table 2 gives an overview of the mean amplitude and standard deviation for the time windows discussed below for the consistent (C) and inconsistent (I) condition, as well as the p-values of paired samples t-tests between the conditions per electrode. 2.1. 300–450 ms In the N400 time window, we found a significant interaction of Location and Consistency, F(2.63, 57.86) = 5.39, p < 0.005 (Greenhouse–Geisser corrected). Follow-up paired samples ttests revealed that the average amplitude in this interval was significantly more negative following inconsistencies than following consistencies at CP1, CP2, P3, P4 and Pz (ps < 0.05), reflecting an increased N400 (which is usually maximal over centroparietal sites, Duncan et al., 2009). 86 BR A IN RE S E A RCH 1 4 18 ( 20 1 1 ) 8 3 –92 Fz (µV) -5 corrected). Follow-up paired samples t-tests revealed that at Fz, FC2 and F4, the average amplitude was more positive for trait inconsistencies than for trait consistencies (ps < 0.01), reflecting an increased LPP. The largest difference was found at Fz. At P3, P4, CP1, CP2 and Pz, the inverse was true, with less positive amplitudes following inconsistencies compared to consistencies (ps < 0.05). 0 LPP 5 0 200 400 600 800 1000 ms 2.3. Temporal PCA Cz (µV) -5 The data was decomposed into 14 temporal factors (TFs) based on inspection of the scree plot, which indicates the gain in explained variance of the grand average after adding more factors (Dien, 2010). Of these factors, there were 8 factors accounting for more than 5% of the variance in the grand averages. Of these 8, in turn, there were 4 that peaked after 250 ms, together accounting for 56% of the variance over the entire 1.2 s trial. Fig. 2 displays the influence of these 4 largest factors on the grand average at Cz. As can be seen, three of the largest factors (TF1, TF3 and TF4) had a positive influence on the amplitude at Cz, and this was true for all thirteen selected channels. TF2 had a negative influence at all thirteen sites. With respect to the negative factor, TF2 was maximal at Pz, peaking at approximately 386 ms post stimulus. Fig. 3 displays the influence of this factor on the grand average for this site as a function of consistency. The negative influence of TF2 proved to be greater for inconsistent than for consistent sentences in the 300–450 ms window at several sites (C3, P3, CP1, Pz, CP2, P4, ps < 0.05). For all selected sites, the average difference between consistent and inconsistent sentences for this factor in the window 300–450 ms correlated significantly with the same difference in the grand average in the window 300–450 ms (mean r = 0.85, ps < 0.001). This indicates that this factor explains the amplitude difference in this time interval to a large extent. We then located the source of this component using sLORETA (Pascual-Marqui, 1999; Pascual-Marqui et al., 1994). Fig. 4 depicts which voxels were maximally involved in eliciting this factor. As can be seen, the left temporo-parietal area, specifically the 0 5 0 200 400 600 800 1000 800 1000 ms -5 Pz (µV) N400 0 5 0 200 400 600 ms Fig. 1 – Grand averages per condition for three midline sites. Grand average waveforms for consistent (full lines) and inconsistent (dotted lines) ending sentences at three midline sites. Negativity is plotted upwards. 2.2. 450–1000 ms Again, we found a significant interaction effect of Location and Consistency, F(2.87, 63.06) = 9.65, p < 0.001 (Greenhouse–Geisser Table 2 – Mean amplitude, standard deviation and p-value of the paired samples t-test per time window for the consistent (C) and inconsistent (I) condition per electrode. 300–450 ms (N400) Fz Cz Pz F3 F4 FC1 FC2 C3 C4 CP1 CP2 P3 P4 Note. ⁎⁎ p < 0.01. ⁎ p < 0.05. 450–1000 ms (LPP) M(C) SD(C) M(I) SD(I) p M(C) SD(C) M(I) SD(I) p −1.43 −0.87 −0.32 −1.53 −1.22 −1.32 −1.11 −0.75 −0.76 −0.35 −0.37 0.64 0.40 2.12 2.28 2.14 1.75 1.77 1.95 1.92 1.59 1.57 1.89 2.09 1.34 1.74 − 0.98 − 1.27 − 1.24 − 1.16 − 1.12 − 1.24 − 1.08 − 1.11 − 1.04 − 1.14 − 1.18 0.05 − 0.17 3.04 2.43 2.49 2.50 2.47 2.69 2.40 1.92 1.99 2.14 2.25 1.40 2.07 0.247 0.202 0.007 ⁎⁎ 0.274 0.776 0.840 0.916 0.117 0.288 0.006 ⁎⁎ 0.009 ⁎⁎ 0.017 ⁎ 0.009 ⁎⁎ 0.81 2.13 2.43 0.22 0.54 1.19 1.03 1.22 1.36 2.31 2.36 1.69 1.64 2.17 2.08 1.70 1.68 1.83 1.98 2.22 1.44 1.46 1.63 1.71 1.24 1.26 1.96 2.07 1.47 0.87 1.26 1.65 1.87 0.92 1.36 1.50 1.61 1.08 1.13 1.99 1.81 1.76 1.53 1.62 1.77 1.97 1.11 1.72 1.51 1.76 1.29 1.38 0.004 ⁎⁎ 0.825 0.007 ⁎⁎ 0.064 0.010 ⁎ 0.189 0.009 ⁎⁎ 0.270 0.997 0.010 ⁎ 0.033 ⁎ 0.011 ⁎ 0.043 ⁎ 87 BR A I N R ES E A RCH 1 4 18 ( 20 1 1 ) 8 3 –92 -1.5 TF2 -1 -0.5 Cz (µV) 0 0.5 1 1.5 TF3 2 TF4 2.5 TF1 3 0 200 400 600 800 1000 Fig. 2 – Projection of the four largest temporal factors to the Cz channel. Projection of the 4 largest temporal factors to the Cz channel. This image is representative for all analyzed channels as far as polarity of the peak of these factors goes. superior temporal gyrus (BA22) was found to be most involved in producing TF2 (MNI coordinates −60, −60, 20). With respect to the positive factors, TF1 was maximal at Pz, while TF3 and TF4 both peaked at Fz. The combined influence of these three factors constituted a significantly larger positivity for inconsistent than for consistent sentences at F3 and Fz (ps < 0.05), while it was smaller for inconsistent sentences at P3, CP1, Pz, CP2 and P4 (p < 0.01–0.05). Fig. 3 shows the combined Fz (µV) -2 -1 0 1 0 200 400 600 800 1000 0 200 400 600 800 1000 0 200 400 600 800 1000 Cz (µV) -2 -1 0 1 Pz (µV) -2 -1 0 1 Fig. 3 – Influence of the four largest temporal factors per condition. Fig. 3 depicts the influence of the four largest temporal factors on three midline channels. The influence of TF 2 is denoted in red, the combined influence of TF 1, 3 and 4 in blue. Full lines correspond to consistent ending sentences, dotted lines to inconsistent endings. Negativity plotted upward. 88 TF 4 TF 3 TF 2 TF 1 BR A IN RE S E A RCH 1 4 18 ( 20 1 1 ) 8 3 –92 Fig. 4 – Source localization of the four largest temporal factors. Estimated current source density for the voxels maximally active in producing the four largest temporal factors. influence of TF1, TF3 and TF4 on three midline channels (Fz, Cz, Pz). Fig. 4 shows the sLORETA source localization of these three components. Voxels in the left cuneus (−10, −100, 20; BA19), medial parietal lobe (5, −45, 71; BA 5) and left cuneus (−30, −95, −5; BA 18) were maximally involved in the generation of TF1, TF3 and TF4 respectively. 3. Discussion The aim of this study was to test whether trait inconsistencies generate increased N400 amplitudes, reflecting violation of expectations based on prior behaviors of others, as well as increased BR A I N R ES E A RCH 1 4 18 ( 20 1 1 ) 8 3 –92 LPPs, presumably reflecting evaluative inconsistency. Both hypotheses were confirmed. 3.1. N400 in spontaneous trait inferences As expected, in the 300–450 ms interval, an increased negative mean amplitude was observed in response to trait inconsistencies at centro-parietal sites. We interpreted this negativity as an N400. This was confirmed by a temporal PCA, which revealed that the second largest temporal factor was a sharp negativity peaking at about 400 ms. This factor had a negative influence on the amplitude at all analyzed electrode locations. The difference between consistency conditions in this time interval could to a great extent be explained by differences in this factor. The neural source that contributed most to this component was located at the left superior temporal gyrus, more specifically in Wernicke's area, which is one of the most consistently reported generators of the N400 (for a review, see Van Petten and Luka, 2006). Taken together, this clearly seems to support an interpretation of the differences in the 300–450 ms as an increased N400 in response to trait inconsistencies. This is in line with previous ERP research on stereotypes (Van Berkum et al., 2008; White et al., 2009). In these studies, violations of stereotypes, another type of person schema, were associated with increased N400 amplitudes. However, no such modulation of the N400 component has been reported before in trait inference studies using paradigms similar to the present research (Bartholow et al., 2001, 2003; Van Duynslaeger et al., 2007, 2008; Van Overwalle et al., 2009). This might be due to differences in the stimulus presentation procedure, as outlined above (see Section 1.2). To repeat briefly, we reasoned that by using multiple critical sentences per scenario, N400 amplitudes in response to trait inconsistencies might have been reduced in the past, as this caused some inconsistencies to be followed by consistencies, and some inconsistencies to be followed by inconsistencies as well, rendering the consistency ambiguous in these cases. Moreover, the presence of a large LPP might have further obscured possible N400 modulations. In line with a classic view of the N400, we propose that the increased amplitude of this component following trait inconsistent behaviors reflects the increased effort required to understand these inconsistent sentences, as they are more difficult to integrate with the previous context (Federmeier and Laszlo, 2009). It seems unlikely that the present modulation of the N400 would be a consequence of evaluative incongruence. Recent research (Holt et al., 2009) has shown that the N400 amplitude following emotional words in a neutral context was larger than following neutral words when participants were explicitly asked to evaluate their emotional content. However, no difference was found between positive and negative words, and an influence of differences in cloze probability could not be entirely excluded. Moreover, a recent study by Herring et al. (2011) directly addressed the possible influence of evaluative incongruence on the N400 and LPP. These authors found no evidence for an influence of evaluative incongruence on the N400. 3.2. Late positivities in spontaneous trait inferences In the later 450–1000 ms interval, increased positive amplitudes were observed at several frontal locations, in line with 89 many previous findings that trait inconsistencies evoke increased LPPs (Bartholow et al., 2001, 2003; Van Duynslaeger et al., 2007, 2008; Van Overwalle et al., 2009). The reversed effect observed at posterior sites could be the result of the use of the average reference, possibly reinforced by the N400 effect, which was larger for inconsistencies at the posterior sites, thus possibly leading to an apparently smaller LPP. In any case, the largest differences were observed at frontal sites. The temporal PCA derived three large late positive factors in the 450–1000 ms time window, all contributing to LPPs in the grand average. In the recent literature, later positivities have been interpreted as reflecting increased attention due to the salience of evaluatively incongruent stimuli. In a recent metaanalysis on the LPP and emotion, Hacjak et al. (2010) concluded that “the LPP reflects multiple and overlapping positivities beginning in the time range of the classic P300, and that these positivities reflect increased salience of the stimuli” (p. 147). Moreover, source localization indicated that the cuneus and medial parietal lobe were most involved in generating these positivities. The important role of these areas in generating the LPP has been confirmed by fMRI research (for a review, see Sabatinelli et al., 2007), and interpreted as a consequence of the motivational relevance of emotional stimuli. To illustrate, it has been demonstrated that spider phobics show increased LPP amplitudes in response to pictures of spiders compared to healthy controls and that both the cuneus and the medial parietal cortex were more engaged by spider pictures in these patients (Scharmüller et al., 2011). However, exactly which cognitive processes are reflected by the increased LPP in response to trait inconsistencies in this and all previous trait studies, cannot be concluded based on the present data. Previous researchers (Bartholow et al., 2001, 2003; Van Duynslaeger et al., 2007, 2008; Van Overwalle et al., 2009) have interpreted it as a P300, reflecting context updating processes. The fact that the increased LPP, unlike the N400, occurs regardless of the number of critical sentences, is compatible with an alternative account, namely that it is a function of evaluative incongruence. It is important to note that the right frontal distribution of the present LPP effect is different than the posterior distribution usually observed in response to evaluative incongruence (Cacioppo et al., 1996) and trait inconsistencies (Bartholow et al., 2003; Van Duynslaeger et al., 2008). There might be several reasons for this discrepancy. First, the difference in distribution could be due to the use of a different reference in the present study. Supporting this notion, Van Duynslaeger et al. (2007), who also employed an average reference like in the present research, found a centrally distributed LPP effect (and no significant differences at the Pz site). Second, Cunningham et al. (2005) showed that the valence of stimuli itself may exert an influence on the distribution of LPP amplitudes, independent of congruence. They found that negative stimuli elicited larger LPPs at right frontal sites than positive stimuli, especially during evaluative judgment (albeit of single, nontrait words). The fact that all our inconsistent sentences were of strong negative valence may therefore be a tentative explanation for the right frontal distribution of the LPP in the present study. Interestingly, in a recent study, Leuthold et al. (2011) found increased N400 and LPP amplitudes in response to verbally 90 BR A IN RE S E A RCH 1 4 18 ( 20 1 1 ) 8 3 –92 described unanticipated emotional reactions (vs. predictable reactions) of unknown actors. Although it must be noted that this study wasn't concerned with trait-based expectations, the LPP reported by Leuthold et al. (2011) had a frontal distribution similar to that in the present research. 3.3. Limitations An important limitation of the present study is that it only included negative violations of positive traits. As such, we are unable to investigate the influence of valence in detail, as this would require the inclusion of positive violations of negative traits. It must be noted here that prior ERP research on spontaneous trait inferences consistently failed to find significant differences between negative consistent and positive inconsistent ending sentences following a negative trait-implying context, either in N400 or LPP amplitude (Bartholow et al., 2003; Van Duynslaeger et al., 2007, 2008). The results of Bartholow et al. (2001) are more difficult to interpret in this respect, because no separate results were reported for negative and positive inconsistencies. As in the present study, Van Overwalle et al. (2009) used only negative inconsistencies. 3.4. General conclusions In sum, we found increases in both N400 and LPP amplitude following behavior descriptions which were inconsistent with a previously implied personality trait. We propose that the increase in N400 amplitude following trait inconsistent behavior descriptions reflects increased effort required to understand these descriptions, as they clearly violate expectations based on prior behaviors of the actor. We interpreted the increased LPP as a consequence of evaluative incongruence. However, based on the present data, it is theoretically not possible to completely rule out that the N400 might also reflect valence evaluation (but see Herring et al., 2011). Further research could address the role of the LPP and N400 in spontaneous trait inferences more precisely by systematically comparing first and second violations of established traits, by keeping the valence of inconsistent behaviors and prior descriptions constant, or by systematically manipulating both valence and inconsistency. In any case, future research aimed at unveiling the cognitive processes underlying trait inferences should take into account that a trait entails both a valuation of past behavior and a prediction about future behavior. 4. Experimental procedures 4.1. Participants 25 students of the Vrije Universiteit Brussel who reported to be right-handed, participated in exchange for course credit. None had a prior history of any neurological dysfunction. The participants comprised of 19 women and 6 men, with an age varying between 18 and 23 (M = 19.15, SD= 1.17). 3 participants made use of reading glasses or lenses during the experiment. 4.2. Stimulus material The ERP-design was a modification of the expectancy-violation paradigm applied by Bartholow et al. (2001, 2003) and Van Duynslaeger et al. (2007, 2008). Participants read 120 sets of Dutch sentences, consisting of 2 or 3 positive trait-implying sentences and one critical sentence. Each sentence consisted of 4 to 7 words, and was presented word by word for 300 ms with an interstimulus interval of 350 ms. Fictional ‘Star Trek’ names were used to avoid association with an existing person (Hoffman and Hurst, 1990). Examples of the trait-implying sentences (translated from Dutch) are: “Diplaq says hello to everybody” and “Diplaq gives his colleague a present” (implying that Diplaq is a friendly person). The last sentence was either trait-consistent (TC) or traitinconsistent (TIC), and the degree of consistency was determined by the last word of the sentence. TC-sentences described a positive behavior that was consistent with the previously implied trait (e.g., “Diplaq gives his mother a hug”). TIC-sentences described a negative behavior that was inconsistent with the previously implied trait. (e.g. “Diplaq gives his mother a slap”). Consistent and inconsistent critical sentences contained the same verbs, and were matched with respect to the amount of words and the number of syllables in the last word. Part of the stimulus material was borrowed from Van Duynslaeger et al. (2007) and additional sentences were developed. All sentences were pilot tested on college students (N = 132) which rated the degree of consistency between each individual sentence and the proposed trait on an 11-point scale ranging from 0 = entirely inconsistent to 10 = entirely consistent. The participants also rated each sentence on valence on an 11point scale going from 0 = very negative to 10= very positive. Sentences with both a mean consistency rating and a mean valence rating higher than 7 were kept as TC-sentences. Sentences with both a mean inconsistency and a valence rating lower than 3 were selected as TIC-sentences. The N400 amplitude following the presentation of words is inversely related to their frequency of use in a given language (Federmeier and Laszlo, 2009). Therefore, we carried out an analysis of frequency of use of the critical end-words, based on the SUBTEX-NL database (Keuleers et al., 2010). This analysis revealed no statistically significant difference in word frequency between conditions (Wilcoxon W= 30932, p = 0.12). The absolute mean frequency was even higher for the inconsistent condition than for the consistent condition, which should, if anything, work against the hypotheses regarding the N400. 4.2.1. Electrophysiological registration and analysis The EEG was recorded from 30 scalp sites according to the international 10–20 electrode system, using sintered AgCl electrodes fixed in a Waveguard cap from Advanced Neuro Technology (ANT). The montage included six midline sites (Fpz, Fz, Cz, Pz, Poz, Oz) and twelve sites over each hemisphere (Fp1/Fp2, F3/ F4, F7/F8, FC1/FC2, FC5/FC6, C3/C4, T7/T8, CP1/CP2, CP5/CP6, P3/P4, P7/P8, O1/O2), with the average of all EEG-channels as recording and off-line reference, as we failed to obtain good mastoid recordings due to technical difficulties. The ground electrode (AFz) was located between the Fz and Cz electrodes. Electro-oculograms (EOGs) were recorded to measure the vertical and horizontal eye movements by means of electrodes placed above and below the right eye and 1 cm external to the BR A I N R ES E A RCH 1 4 18 ( 20 1 1 ) 8 3 –92 outer canthus of each eye, respectively. Impedance was kept below 10 kΩ for each electrode. The EEG and EOG were continuously recorded during the experiment at a digitizing rate of 256 Hz with a DC amplifier. An offline 0.01- to 30-Hz 4th order Butterworth band pass was applied (Bartholow et al., 2001; 2003). Epochs were extracted, composed of 1000 ms following the onset of the last word of critical sentences as well as the 200 ms preceding it, which served as sample-period for baseline-correction. Epochs containing ocular or muscular artifacts were removed, based on visual inspection of the data. Two participants were excluded, based on the criterion of more than 33% rejected trials. For the remaining 23 participants, on average 20% of the trials were rejected, resulting in 48 valid trials per condition on average. The stimuli were presented on a TFT-screen with a refresh rate of 25 ms at a distance of 50 cm. Stimuli were presented with E-prime, from Psychology Software Tools, Incorporated. EEG was recorded and processed with hardware (Cognitrace) and software (ASA, Eemagine) developed by ANT. 4.3. Procedure Participants sat down in a comfortable chair, received information about the procedure, and signed the informed consent, which was approved by the medical ethical committee of the Vrije Universiteit Brussel. They were informed that they would read sentences about a protagonist's behavior which would be presented word by word on a computer screen. Every participant was instructed to read the material attentively and to “try to familiarize yourself with the material of the experiment,” which is a typical instruction to elicit spontaneous trait inferences (see Todorov and Uleman, 2002). The participants received four practice runs prior to the actual experiment. 4.4. 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