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Available online at www.sciencedirect.com

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Research Report

What is the specificity of the response to the own first-name when presented as a novel in a passive oddball paradigm? An ERP study Jean-Baptiste Eichenlaub⁎, Perrine Ruby1 , Dominique Morlet1 INSERM U1028, Lyon Neuroscience Research Center, Brain Dynamics and Cognition Team, Lyon, F-69000, France CNRS UMR5292, Lyon Neuroscience Research Center, Brain Dynamics and Cognition Team, Lyon, F-69000, France University Lyon 1, Lyon, F-69000, France

A R T I C LE I N FO

AB S T R A C T

Article history:

One's own first-name is a special stimulus: one's attention is more likely captured by hear-

Accepted 29 January 2012

ing one's own first-name than by hearing another first-name. Previous event-related poten-

Available online 8 February 2012

tial (ERP) studies demonstrated that this special stimulus produces differential responses both in active and in passive condition. Such results suggest that passively hearing one's

Keywords:

own first-name triggers processing levels generally activated by the explicit detection of

Self

stimuli. This questions about the particular power of the own first-name to automatically

Familiarity

orient attention, but no study investigated the specific response to this special stimulus

Attention orienting

in a paradigm designed to study automatic attention orienting. In this ERP study, we com-

Event-related potential

pared the responses elicited by the own first-name (OWN) and one unfamiliar first-name

Novelty P3

(OTHER) presented, rarely, randomly and at the same frequency among repetitive tones (i.e. as novel stimuli in an oddball paradigm) while subjects (N = 36) were watching a silent movie with subtitles. We tested at what latency the responses to OWN and OTHER diverge, and whether OWN modulates the brain orienting response (novelty P3). Data analysis showed specific responses to OWN after 300 ms. OWN only evoked a central negativity (320 ms) and a parietal positivity (550 ms). However, OWN had no significant effect on the brain orienting response (260 ms). Our results confirm that the own first-name does elicit a late specific brain response. However, they challenge the idea that in passive condition, the own first-name is systematically more powerful than another first-name to orient attention when it is heard unexpectedly. © 2012 Elsevier B.V. All rights reserved.

1.

Introduction

One's own first-name is a special stimulus. It refers to oneself, it is heard since birth, it is one of the predominant means used

by others to engage in social interactions and it is mainly used by related (closely or not) people. As a consequence, hearing one's own first-name and hearing another first-name yield different behavioral responses. For instance, everyone cer-

⁎ Corresponding author at: INSERM U1028, CNRS UMR5292, Centre de Recherche en Neurosciences de Lyon, Equipe Dynamique Cérébrale et Cognition, Centre Hospitalier le Vinatier, Bâtiment 452, 95 Bd Pinel, Bron, F-69500, France. Fax: +33 4 72 13 89 01. E-mail address: [email protected] (J.-B. Eichenlaub). 1 Contributed equally. 0006-8993/$ – see front matter © 2012 Elsevier B.V. All rights reserved. doi:10.1016/j.brainres.2012.01.072

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tainly noticed that the utterance of one's first-name in the room very often shifts one's attention even though one is engaged in a discussion or another ongoing task. Experimental psychology studies measured this effect and confirmed that the own first-name is particularly prone to capture attention even when one is involved in an auditory task. Two studies showed that the own first-name is more powerful than another word or first-name to involuntarily capture attention when presented in the left ignored ear while the subject listens and shadows simultaneously a discourse presented in the right ear (Moray, 1959; Wood and Cowan, 1995). In the same line, a study combining behavioral and electrophysiological assessment (through skin conductance and cardiac activity) showed that the written own first-name presented as an unexpected stimulus elicited an enhanced orienting response as compared to control stimuli (Shek and Spinks, 1985). A way to better understand this “own fist-name effect” is to investigate the brain responses to the subject's first-name. Several studies assessed the auditory event-related potentials (ERPs) elicited by the subject's first-name using various paradigms (Berlad and Pratt, 1995; Folmer and Yingling, 1997; Holeckova et al., 2006; Holeckova et al., 2008; Muller and Kutas, 1996; Perrin et al., 1999; Perrin et al., 2005; Perrin et al., 2006; Pratt et al., 1999). Most of them demonstrated that this special stimulus produces positive responses at latencies from around 300 ms after stimulus onset, that were larger than the responses evoked by common nouns (Berlad and Pratt, 1995; Pratt et al., 1999), other first-names (Folmer and Yingling, 1997; Muller and Kutas, 1996; Perrin et al., 1999; Perrin et al., 2005; Perrin et al., 2006) or non-vocal stimuli (Holeckova et al., 2006). Such a result was obtained whenever subjects paid attention to their first-name (Perrin et al., 1999), did not pay attention to the stimuli (Berlad and Pratt, 1995; Folmer and Yingling, 1997; Muller and Kutas, 1996; Perrin et al., 1999; Perrin et al., 2005; Perrin et al., 2006; Pratt et al., 1999) or were distracted from the stimuli by a visual task (Holeckova et al., 2006; Pratt et al., 1999). Noteworthy, the subject's first-name can still elicit a differential brain response when the subject is asleep (Perrin et al., 1999; Pratt et al., 1999) or in a pathological state of altered consciousness like coma or vegetative state (e.g. Fischer et al., 2008; Fischer et al., 2010; Perrin et al., 2006; Schnakers et al., 2008). These results led some authors to suggest that the subject's first-name is “automatically and implicitly processed as a target stimulus” (Perrin et al., 1999). The fact that the own first-name can trigger the processing levels that are generally activated by target stimuli (i.e. stimuli that subjects have to detect explicitly) even when subjects/ patients do not pay attention to it, questions about the specific power of this relevant stimulus to automatically orient the attention. Automatic attention orienting brain responses to auditory stimuli can be assessed through so-called oddball novelty paradigms. In such paradigms, an unexpected and salient stimulus (novel) is presented randomly and rarely among repetitive tones. An extensive literature studied the capture of attention by novel stimuli of different types, in various experimental conditions (Friedman et al., 2001; Polich, 2007; Ranganath and Rainer, 2003). Only one ERP study (Holeckova et al., 2006) used a novelty oddball paradigm with the own first-name as novel. However,

this study was designed to investigate the brain response to a familiar voice. It is not possible from Holeckova's results to disentangle a specific effect of the own first-name from a more general effect of verbal stimuli. Thus, while the own first-name is known to easily capture attention, the actual specificity of the brain response to the own first-name presented as a novel was never assessed. In order to assess the specificity of the response to the own first-name, we compared it to the response to a control stimulus. In order to minimize the confounding factors we selected and presented the control stimulus as explained below. Firstly, much more levels of language processing are controlled if the control stimulus and the own first-name belong to the same lexical/semantic category. Using non vocal stimuli (Holeckova et al., 2006) or common nouns (Berlad and Pratt, 1995; Pratt et al., 1999) as control stimuli may induce some lexical/semantic differential processing that would interfere with the specific effect of the own first-name. We used another first-name as control stimulus to minimize this bias (Folmer and Yingling, 1997; Muller and Kutas, 1996; Perrin et al., 1999; Perrin et al., 2005; Perrin et al., 2006). Secondly, acoustic differences between the own first-name and the control stimulus cannot be suppressed. Nonetheless, matching the attacks and envelops of the two sounds avoid modulation of the sensory responses (and then possibly of the subsequent responses) by these parameters (Näätänen and Picton, 1987). Finally, as the brain response is modulated by the frequency of presentation of a stimulus, matching the frequency of presentation of the own first-name and the control stimulus prevent from confounds. In some previous studies (Perrin et al., 1999; Perrin et al., 2005; Perrin et al., 2006), the own first-name and several other first-names were equiprobably presented and the response to the own firstname was compared with the averaged responses to the other first-names. If each first-name had the same probability of presentation, nonetheless the control stimuli i.e. irrelevant names, were more frequently presented than the relevant stimulus i.e. the own first-name. In such conditions, an enhanced brain response to the own first-name could be due not only to the self-reference of the stimulus, but also to the rarity of presentation of this relevant stimulus as compared with the irrelevant ones (Polich, 2007). As a consequence such a paradigm was not optimum to assess the specificity of the response to the own first-name. In the present study, to investigate the specificity of the response to the own first-name, we used as control stimulus one first-name (to control the contrast of familiarity between the two first-names, first-names belonging to persons in the close environment of the subjects were excluded). We matched as much as possible the attacks and the envelops of the two stimuli and we presented each of them at the same frequency as novels in an auditory oddball novelty paradigm. Novels typically evoke a fronto-central positive component between 220 and 320 ms, the so-called novelty P3 (nP3) (Friedman et al., 2001; Ranganath and Rainer, 2003). This component, which correlates with autonomic manifestations of the orienting response (Barry et al., 2010; Knight, 1996; Lyytinen et al., 1992; Marinkovic et al., 2001), is interpreted as reflecting the orienting of attention towards the unexpected stimulus, hence commonly referred to as the “brain orienting response” (Friedman et al., 2001; Ranganath and Rainer, 2003).

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It was suggested that the larger the novelty P3 the stronger the orientation of attention (Dominguez-Borras et al., 2008; Escera et al., 2003). As behavioral studies showed that the own firstname more easily orients attention than any other names (Moray, 1959; Wood and Cowan, 1995), we expected a larger novelty P3 in response to the own first-name than to the other first-name. At later latencies (around 500 ms) a novel sound can also elicit parietal components that may be related to familiarity (Holeckova et al., 2006), semantic (Mecklinger et al., 1997) or memory processing (Friedman and Simpson, 1994; Holeckova et al., 2006). We thus expected that the own first-name, which is familiar and associated with many episodic memories, would elicit larger late responses than the unfamiliar first-name. The objective of this study was to investigate the specificity of the response to the own first-name in a context of automatic attention orienting. We used a novelty oddball paradigm in which the own first-name (OWN) and one other first-name (OTHER), were presented rarely, randomly and at equal frequencies (p = 0.02 each) among repetitive pure tones. During ERPs acquisition, the participants (N = 36) were watching a silent self-selected movie with subtitles. We compared the time course and morphology of the brain responses elicited by OWN and OTHER to test at which latency the two responses diverge and whether the own first-name modulates the brain orienting response.

2.

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Results

The responses to OWN and to OTHER averaged over 36 subjects are displayed in Fig. 1. The number of accepted trials for each subject was not different between OWN and OTHER (paired t-test not significant). It was in average 77 ± 18 (63% of presented trials ± 15) for OWN and 78 ± 18 (65% ± 14) for OTHER. Both novels elicited the auditory N1, a novelty P3 (nP3) and a large temporal positive component (TP). OWN only evoked a short central negative component (CN) and a large and long-lasting parietal positive response (PP).

2.1.

Components evoked both by OWN and OTHER

2.1.1.

Auditory N1

The auditory N1s evoked by OWN and by OTHER were not different in the 140–180 ms interval (Figs. 1 and 2). They peaked both around 160 ms at Cz (see Table 1). On scalp potential (SP) maps, the N1 showed as expected a negative centrally distributed topography. This response pattern is typical of activity in both auditory cortices, and appeared in the scalp current density (SCD) maps as strong current sinks over bilateral centrolateral regions. No difference was observed between the maps (SP and SCD, Fig. 3) elicited by OWN and by OTHER in the 140–180 ms time interval.

Fig. 1 – Grand average waveforms recorded at 15 scalp electrodes (among 21 recorded electrodes) and at 2 peri-ocular electrodes (EOG1 at the supraorbital ridge of the left eye and EOG2 at the infraorbital ridge of the right eye) for the 36 subjects, in response to the own first-name (OWN) and to the unfamiliar first-name (OTHER), both presented as novels (p = 0.02) in an auditory passive oddball paradigm. The blue-colored areas on the ERPs show the electrodes on the time-intervals of interest (i.e. 300–340 ms for Central Negativity and 450–650 ms for Parietal Positivity) in which discrepancies between the responses to OWN and OTHER were significant. The stars indicate the result of Wilcoxon matched rank sign test comparing mean amplitudes of the responses to OWN and OTHER. *p < 0.05, **p < 0.01, ***p < 0.001. At EOG1 and EOG2, no significant difference was found between OWN and OTHER.

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Fig. 2 – Statistical significance of Wilcoxon matched rank sign test (p < 0.05) performed at each sampling point and at 15 electrodes assessing: (1) Evoked responses versus baseline (OWN in red, OTHER in black), (2) Response evoked by OWN versus response evoked by OTHER (in gray).

2.1.2.

Novelty P3

In the 230–280 ms time-interval, OWN and OTHER elicited not different but significantly positive waves (see Figs. 1 and 2) referred to as “novelty P3” (Friedman et al., 2001; see Fig. 3). They peaked both around 260 ms at Cz at non-significantly different latencies (see Table 1). SP maps in the 230–280 ms time

interval (Figs. 3 and 5) showed a positive fronto-centrally distributed topography. SCD maps in the same time-window (Fig. 3) revealed positive currents over central locations. No difference was observed between the maps (SP and SCD) elicited by OWN and by OTHER. Novelty P3 peak amplitudes were also not different (Table 1) and this result of a non

Table 1 – Mean amplitudes (Mean) ± SD (in μV), peak amplitudes (Max) ± SD (in μV) and peak latencies (Lat) ± SD (in ms) of the responses to the own first-name (OWN) and to the unfamiliar first-name (OTHER) over the 5 intervals of interests: 140–180 ms at Cz (N1), 230–280 ms at Cz (novelty P3), 300–340 ms at Cz (CN), 250–450 ms at T4 (TP) and 450–650 ms at Pz (PP). CN and PP being not detected in response to OTHER, their peak amplitude and peak latency were not measured (noted Ø). The last column shows the result of the Wilcoxon matched rank sign test comparing the peak amplitudes, peak latencies and mean amplitudes of the responses to OWN with those of the responses to OTHER. *p < 0.05;***p < 0.001; ns, non-significant; Ø, non appropriate.

N1 Cz; 140–180 ms

Novelty P3 Cz; 230–280 ms

CN Cz; 300–340 ms

TP T4; 250–450 ms

PP Pz; 450–650 ms

Mean (μV) Max (μV) Lat (ms)

OWN

OTHER

OWN vs OTHER

− 4.1 ± 3.4 − 5.3 ± 3.8 161 ± 14

−4.1 ± 3.0 −5.3 ± 3.3 160 ± 15

ns ns ns

Mean Max Lat

2.6 ± 4.0 4.4 ± 4.6 260 ± 16

2.6 ± 4.0 4.2 ± 4.2 263 ± 16

ns ns ns

Mean Max Lat

− 0.1 ± 2.9 − 1.3 ± 2.8 328 ± 13

1.2 ± 3.8 Ø Ø

* Ø Ø

Mean Max Lat

2.2 ± 1.6 4.2 ± 1.9 355 ± 75

1.8 ± 1.6 3.6 ± 1.7 337 ± 61

ns ns ns

Mean Max Lat

2.8 ± 3.0 5.4 ± 3.2 547 ± 73

0.5 ± 2.0 Ø Ø

*** Ø Ø

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Fig. 3 – Mean scalp potential (SP) and scalp current density (SCD) distributions of the responses to the own first-name (OWN) and to the unfamiliar first-name (OTHER) over the 140–180 ms (N1), 230–280 ms (novelty P3) and 250–450 ms (TP) time-windows.

different novelty P3 response to OWN and to OTHER was further confirmed in a supplementary analysis in which the mean amplitudes of the ERPs elicited by OWN and OTHER were calculated in a 50 ms time-interval around the individual peak latencies of the nP3 (see Supplementary data).

2.1.3.

Temporal positivity (TP)

In the 250–450 ms time-interval, OWN and OTHER elicited not different but significantly positive waves (Figs. 1 and 2) peaking around 350 ms at T4 (see Table 1). SP maps (Figs. 3 and 5) showed a positive temporally distributed topography. SCD maps (Fig. 3) revealed positive currents over temporal locations. No difference was observed between the maps (SP and SCD) elicited by OWN and OTHER in the 250–450 ms time interval.

2.2.

Components evoked only by OWN

2.2.1.

Central Negativity (CN)

In the 300–340 ms time-interval (Figs. 1 and 2), the response evoked by OWN differed from the response evoked by OTHER. OWN evoked a significantly negative wave (Fig. 2). By contrast, OTHER evoked no significant negative response around this latency. The mean amplitude at Cz within this latency range was significantly more negative for OWN than for OTHER (p < 0.05; Table 1). In SP maps drawn from the data in the 300–340 ms interval (Fig. 4), OWN and OTHER showed a positive temporally distributed topography (corresponding to the Temporal Positive component – TP – described just above), but only OWN

Fig. 4 – Mean scalp potential (SP) and scalp current density (SCD) distributions of the responses to the own first-name (OWN) and to the unfamiliar first-name (OTHER) over the 300–340 ms (CN) and 450–650 ms (PP) time-windows. The third row shows the maps of the differences (OWN minus OTHER). The statistical significance of the comparison (Wilcoxon matched rank sign test) assessed at each electrode is shown with black points at the electrode sites.

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showed an additional negative centrally distributed topography. In the difference distribution (OWN minus OTHER) the temporal positive activity common to OWN and OTHER was removed and only the central distribution of a negative component specific to OWN appeared (Fig. 4). Visual SCD analysis (Fig. 4) revealed similar current density distribution for OWN and for OTHER, with positive sources in temporal region and negative sinks at frontal and parietooccipital locations. However, statistical analysis revealed that the current sinks in the left, middle and right central regions were stronger for OWN than for OTHER.

2.2.2.

Parietal positivity (PP)

At later latencies, the response evoked by OWN largely differed from the response evoked by OTHER (Figs. 1 and 2). OWN evoked a large long-lasting positive wave around 550 ms referred to as parietal positivity (PP), with maximal amplitude at Pz. By contrast, OTHER evoked no significant response around this latency. As a consequence, the mean amplitude at Pz within this latency range was significantly different for OWN and for OTHER (p < 0.001; Table 1). In SP maps (Figs. 4 and 5), OWN, but not OTHER, showed a positive centro-parietally distributed topography and significant differences between the two responses were found on a large number of central and parietal electrodes. For OWN and for OTHER, SCD maps (Fig. 4) showed a positive source in parietal region and a negative sink in central location. The statistical analysis revealed stronger positive source in left parietal region and in occipital location for OWN than for OTHER.

3.

Discussion

The present study aimed at investigating the specificity of the brain response to the own first-name in a context of automatic attention orienting. In 36 healthy subjects, we compared the brain responses to the hearing of the own first-name (OWN) and to the hearing of one unfamiliar first-name (OTHER). First-names were presented at equal low frequencies (as novels) in a passive auditory oddball paradigm i.e. while subjects were watching a silent self-selected movie with subtiltles. Our objective was to compare the time course of the brain responses elicited by the two first-names to assess at which latency the responses to OWN and OTHER diverge and whether OWN modulates the brain orienting response (novelty P3). Each novel stimulus elicited an auditory N1, followed by a central positivity around 260 ms, that we called novelty P3 in keeping with the literature showing a modality non-specific positive response at these latencies to rare, unexpected and salient stimuli (Friedman et al., 2001; Polich, 2007; Ranganath and Rainer, 2003). Unexpectedly, no specific effect of OWN was observed at the latency of the novelty P3. However, the brain response to OWN differed from the brain response to OTHER after 300 ms: a central negativity around 320 ms (CN) and a long lasting (450–650 ms) parietal positivity (PP) were found in response to OWN, but not in response to OTHER. It is to note that OWN and OTHER elicited both a long lasting (250–450 ms) right lateralized temporal positive component (TP).

Fig. 5 – Mean scalp potential (SP) distributions of the responses to the own first-name (OWN) and to the unfamiliar first-name (OTHER) from 250 to 650 ms. These right views highlight the time course of 1) a fronto-central novelty P3 from 240 to 300 ms in response to both first-names, 2) a right temporal positivity from 230 to 450 ms for both first-names and 3) a late parietal positivity from 400 to 720 ms for the own first-name only.

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3.1.

No effect of OWN on the novelty P3

A novel presented in an auditory oddball paradigm typically evokes a novelty P3, a positive component around 260 ms generally showing a frontal–central distribution and presenting common features with the P3a component evoked by deviant stimuli in passive paradigms (Friedman et al., 2001; Herrmann and Knight, 2001; Polich, 2007; Ranganath and Rainer, 2003). This frontal–central novelty P3 (novelty P3a) is interpreted as reflecting the involuntary orienting of attention towards unexpected stimuli (Friedman et al., 2001; Ranganath and Rainer, 2003). Some authors recently proposed that the novelty P3a could be parsed into two separate subcomponents, revealing different stages of the attention orienting (e.g. Brinkman and Stauder, 2008; Escera et al., 1998; McDonald et al., 2010; SanMiguel et al., 2010; Wetzel and Schroger, 2007; Yago et al., 2003). The early novelty P3a (eP3a) is maximal centrally and inverts in polarity at posterior and lateral electrodes while the late novelty P3a (lP3a) is maximal frontally without inversion of polarity at posterior and lateral electrodes (Escera et al., 1998). Furthermore, SCD analysis revealed that a central source contributed largely to the eP3a while frontal and parietal sources contributed to the lP3a (Yago et al., 2003). According to Escera et al. (1998), the eP3a is unaffected by attention manipulation, while the lP3a is enhanced by attention. Although the functional significance of these two parts of the novelty P3a is not clear yet, it was suggested that the eP3a could reflect an alerting process governing the direction of the attentional move (Ceponiene et al., 2004). The novelty P3 may also present at later latencies a parietal subcomponent showing features in common with the P3b elicited by target stimuli (Gaeta et al., 2003; Herrmann and Knight, 2001; Polich and Criado, 2006; Polich, 2007; Ranganath and Rainer, 2003). This parietal subcomponent (novelty P3b) seems to appear when attention towards the stimulus increases (e.g. Holeckova et al., 2006; Polich, 2007). Depending on the authors, the P3b was associated with various cognitive processes such as, for example, stimulus categorization, context-updating operations and subsequent memory storage (Friedman et al., 2001; Gaeta et al., 2003; Sutton et al., 1965). At the behavioral level, the own first-name orients attention more easily than any other name (Moray, 1959; Shek and Spinks, 1985; Wood and Cowan, 1995). At the electrophysiological level, it was suggested that the larger the novelty P3 the stronger the orientation of attention (Dominguez-Borras et al., 2008; Escera et al., 2003). Some studies also suggested that even if a subject is involved in a distractive task, a P3b-like can be elicited if subject's attention is strongly attracted by the novels (e.g., Holeckova et al., 2006). We thus expected that OWN would elicit a larger novelty P3 than OTHER and that at least OWN would elicit a novelty P3 including a parietal subcomponent. However, our data did not confirm our hypothesis i.e. they showed that both stimuli elicited similar novelty P3s without any parietal component. In our study, OWN and OTHER elicited a response showing a pattern typical of an eP3a (a maximum amplitude at Cz and inverted polarity at mastoids in potentials maps and a central current source in SCD maps, see Figs. 1, 2 and 3). An lP3a component could neither be detected for OWN nor for OTHER. This pattern suggests that the two novels triggered alerting

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processes, but that none of them was able to induce the subsequent steps of the attention orienting process. In other words our data show that OWN did not influence the brain orienting response, and more particularly, OWN did not force attention orienting better than OTHER. It is unlikely that the absence of modulation of novelty P3 by OWN in our data could be explained by an inability of the brain to categorize auditory stimuli before 250 ms. First, although OWN and OTHER began with the same phone, they were perceived as different from the beginning of their utterance: they were different from the first syllable, and their first phonemes were uttered differently through the coarticulation phenomenon (Hardcastle and Hewlett, 1999). Second, previous studies in the auditory modality demonstrated that stimulus category (such as identifiability or self-relevance) could modulate the novelty P3 amplitude. This effect was mainly observed when subjects direct their attention towards auditory stimulation (Escera et al., 2003; Mecklinger et al., 1997; Opitz et al., 1999), but it was also reported when subjects are involved in a distractive task (Holeckova et al., 2006; Wetzel et al., 2011). Furthermore, using a two-deviant passive oddball paradigm, Roye et al. (2007; 2010) demonstrated that personal and unfamiliar ringtones evoke ERPs and oscillatory responses diverging before the latency of the P3a. We conclude from these results that it is possible for the brain to categorize complex sounds at early latencies (around 250 ms) in passive condition. Rather, it seems likely that the amount of attentional resources left available for the auditory processing could explain our result. Indeed, attention is a factor known to strongly modulate the novelty P3. Studies manipulating attention in auditory oddball paradigms demonstrated that the more the attention is directed towards the sounds the larger the novelty P3 and the more the P3b component appears (e.g. Friedman et al., 1998; Mecklinger et al., 1997; Opitz et al., 1999). Coherently, it was shown that increasing visual perceptual load and working memory load leads to a decrease of the auditory novelty P3 amplitude (e.g. Berti and Schroger, 2003; Lv et al., 2010; Miller et al., 2011; SanMiguel et al., 2008; Zhang et al., 2006). Attention is also a key parameter in the modulation of novelty processing by novel stimulus category. When subjects have to detect targets and thus to attend the stimuli (active condition), novelty P3s elicited by unexpected stimuli are enhanced when the novel stimuli can be identifiable or are self-related. In the visual modality, the brain response to novelty is modulated by the own face presented as a novel in an active task dealing with famous faces (Ninomiya et al., 1998) and by the written own first-name presented as a novel in an active task dealing with other written characters (Zhao et al., 2011). In the auditory modality, Mecklinger et al. (1997) and Opitz et al. (1999) showed that identifiability of novel sounds could modulate the potentials in the time-window of novelty P3 when the subjects were counting deviant sounds. By contrast, when subject's attention is diverted from the stimuli (passive condition), several studies showed that the novels' category does not modulate the novelty P3 (Escera et al., 2003; Mecklinger et al., 1997; Opitz et al., 1999). Escera et al. (2003) demonstrated that semantic analysis of irrelevant sounds depends on the top-down cognitive influences of the attentional set. In our experimental conditions (unattended auditory stimuli while the subjects are watching a silent move) the absence of

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modulation of novelty P3 by the own first-name, although unexpected in the frame of the “own name” literature, is in agreement with this “novelty” literature (Escera et al., 2003; Mecklinger et al., 1997; Opitz et al., 1999), which suggests that if an auditory distractor is presented outside the attentional set, the brain gives priority to novelty detection whatever the content of the novel. To our knowledge, only two studies found a modulation of novelty P3 by novel category in passive condition (Holeckova et al., 2006; Wetzel et al., 2011). Wetzel et al. (2011) observed an effect of novels' identifiability at the latency of P2/early P3a (170–230 ms). Their paradigm shows substantial differences with our paradigm: they used 96 different novel environmental sounds (48 were identifiable sounds) while we presented only two vocal novels (OWN and OTHER). In their paradigm, the constant renewing of the novels might have differently solicited the alerting processes underlying the early P3 than in our paradigm, thus allowing a modulation by novel's category at these latencies, not observed in our data. Holeckova et al. (2006) found a larger novelty P3 to own first-names than to non-vocal stimuli with matching acoustic characteristics. Interestingly, despite apparently very similar paradigms, our data (no lP3a and no P3b in response to OWN) are strongly different from those of Holeckova et al. (lP3a and P3b in response to the own first-names) (Holeckova et al., 2006), suggesting different amount of attentional resources available for the auditory processing between the two studies. Several underestimated differences in experimental condition could explain this attentional effect. In the first place, the distractive task, though apparently similar (watching a silenced movie), might have been in fact quite different. Holeckova's distractive task was an imposed documentary without subtitles, while our distractive task was a self-chosen fulllength film with subtitles. Our task did strongly capture the subjects' attention as demonstrated by the subject's interview at the end of the recording session (all of them told that they enjoyed the movie and were able to report about the story, and none of them complained about any inconvenience caused by the auditory stimuli). This may have been less the case in Holeckova's study: the moderately absorbing film may have made it possible for the subjects to allow some attention towards the sounds. Furthermore, in our study, both performing the task (reading subtitles) and processing the novels (spoken words) activated language processing and thus recruited at least partly the same cerebral resources (Holeckova et al., 2008). As a consequence, compared with Holeckova's task (Holeckova et al., 2006) (movie without subtitles), our task may have induced a delay in the processing of the content of the linguistic novels. Finally, in Holeckova's study recordings were performed during the day while our recordings were performed at night (starting at 10.24 pm± 46 min). The time of day is known to affect some cognitive performance as well as subjective alertness (e.g. Monk et al., 1997). It may then possibly affect the processing of novelty too. In our study, attention orienting processes are not modulated by the novels' category, even if novel stimuli as relevant for the subject as his/her own first-name are presented. This effect may be explained by the fact that our experimental conditions have particularly enhanced the power of the primary task to capture attention, which probably attenuated the

possibility for the novels to be categorized. We learn from our results that even if the own first-name may have some advantage to orient attention in many conditions, it is not necessarily the case in every experimental passive condition. When the subject's attention is strongly involved in a primary task, at the latency of the novelty P3, priority seems to be given to novelty detection rather than to novels categorization, even if the novel is the own first-name.

3.2.

Specific effect of OWN on late components

The responses to OWN and to OTHER differed after the novelty P3 i.e. after 300 ms. Only OWN elicited a central negative component around 320 ms (CN) and a centro-parietal positive component around 550 ms (PP). Similar negative components in response to novel sounds have been described both in active and in passive conditions (Escera et al., 2001; Holeckova et al., 2006; Holeckova et al., 2008; Kushnerenko et al., 2007; Mecklinger et al., 1997; Opitz et al., 1999; Schroger and Wolff, 1998). However, the cognitive processes subserving such components are not yet fully understood. The CN could reflect the reallocation of attention back to the task (Escera et al., 2001; Kushnerenko et al., 2007; Schroger and Wolff, 1998). It could also reflect further processing of the novels (Escera et al., 2001), as for example conceptual semantic integration processing, i.e. the identification of the mental concept expressed by the novels (Mecklinger et al., 1997; Opitz et al., 1999), or familiarity processing (Holeckova et al., 2006). The first interpretation does not explain our results very well: as the brain orienting response (novelty P3) is of similar amplitude for OWN and for OTHER it seems unlikely that reallocation of attention back to the movie would be more necessary for OWN than for OTHER. The second hypothesis may be more likely: by contrast with OTHER, OWN is heard since birth and is then much more familiar than OTHER. OWN and OTHER have also different meanings for the subjects and much more representations are associated with OWN than with OTHER. The negative component in response to OWN may thus result from either familiarity processing or from conceptual semantic integration processing or from both. In our study, OWN only elicited a centro-parietal positive component around 550 ms. Several previous studies observed a centro-parietal positive component at these latencies in response to the own first-name. This result was consistently obtained despite very different auditory paradigms (Berlad and Pratt, 1995; Folmer and Yingling, 1997; Holeckova et al., 2006; Perrin et al., 1999; Perrin et al., 2005; Perrin et al., 2006; Pratt et al., 1999). It is important to note that the different authors attributed various labels to this late positive component (P3, P300, P450, PP…). Interestingly, Holeckova et al. (2006) clearly dissociated a late parietal positive component (PP) from a novelty P3b component using SCD maps. The SCD maps of the novelty P3b show positive current sources over large frontal and parietal sites (see also Yago et al., 2003), whereas the SCD maps of the PP show negative current sinks over frontal sites and positive current sources over parietal sites. The SCD maps of the parietal positive wave detected in our study (Fig. 4) are similar to the SCD maps of the PP described by Holeckova et al. (2006). As a consequence we concluded that

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the late positive component detected in the present study most probably belonged to the PP family. In the study of Holeckova et al. (2006), PP was obtained in response to the own first-name only when it was uttered by a familiar voice. The PP wave could thus be interpreted as a response to the familiarity of the voice. Our result rules out this explanation and demonstrates that a PP can be obtained in response to the own first-name even if it is not uttered by a familiar voice. As a consequence neither the own firstname nor the familiarity of the voice can explain alone the PP component. The common feature shared by the stimuli that elicited a PP in Holeckova et al. (2006) and in our study may be their significance for the subject. Indeed in both studies, the novel which elicits a PP may be considered as the more meaningful for the subject i.e. one's own name uttered by a familiar voice versus an unfamiliar voice (in Holeckova's study), and the own first-name versus an unfamiliar first-name (in the present study). The novel the more meaningful for the subject may thus induce more semantic and/or recollection processing that the other one. This interpretation of the PP is coherent with previous ERPs studies, which associated a parietal positive wave with semantic processing and/or with the retrieval of information from memory (e.g., Curran, 2004; Friedman and Simpson, 1994) or with emotional processing (e.g., Kissler et al., 2009). The design of our study alone did not make possible further characterization of the late cognitive processes evoked by OWN only. In the light of our results and of other results from the literature, we can only suggest that these late processes were elicited by the own first-name because of its specific importance for the subject relative to the other first-name. In the specific context of our experiment, they might be related to features that distinguish OWN from OTHER: self-reference (i.e. OWN designates the self), emotional content (e.g. OWN is used by related people), familiarity (i.e. OWN is repetitively heard during the whole life) and a number of associated representations (e.g. OWN is associated with a lot of episodic memories whereas OTHER is associated with very few memories).

3.3. sites

A similar response to OWN and to OTHER at temporal

Both OWN and OTHER elicited a similar long lasting (250–450 ms) right lateralized temporal positive component (TP). This late component presents several common features with the fronto-temporal positivity to voice (FTPV) described by Rogier et al. (2010) in children and by Charest et al. (2009) in young adults. It is possible that, in our study, this positivity reflects a response to voice. In this case, as all the stimuli were uttered by the same voice, it makes sense that OWN and OTHER elicited a similar TP.

making difficult the comparison of results from the literature. An alternative way to objectively assess to what extend the attention is effectively kept away from the auditory stimuli would be the use of a high demanding visual continuous performance task (e.g. Escera et al., 2003). Along the same line, another limitation of this study is the impossibility to get a direct behavioral measure of the attentional capture by novels. This is an intrinsic limitation of passive paradigms. As a consequence, in order to confirm our interpretation of the results, further studies controlling the alertness of the subjects and using a high demanding visual continuous performance task and/or measuring attentional capture at the behavioral level are needed. Finally, in this study, the late effects (i.e. the responses elicited only by OWN after 300 ms) are difficult to interpret in terms of precise cognitive processes. For instance, even if we selected OTHER so that it was not familiar to the subject, we did not control for the frequency of use of OWN and OTHER. As a consequence the effect of familiarity cannot be precisely assessed in this study.

3.5.

Conclusion

Our paradigm made it possible to assess the time course of the brain responses specifically elicited by the own first-name versus an unfamiliar first-name in a context of passive automatic attention orienting. When presented rarely and unexpectedly (as novel) while the subject is involved in an absorbing visual task (in passive condition), the own firstname does not modulate the brain orienting response (novelty P3). From this result we suggest that, when a subject is involved in a task, his/her own first-name does not systematically have an advantage to automatically orient his/her attention. Specific components were elicited by the own first-name after 300 ms: a central negative component and a long-lasting parietal positive component. This result suggests that despite its absence of advantage in attention orienting, the own first-name can induce some complex cognitive processing, not necessarily conscious, such as memory recall, or emotional processing, that another, irrelevant, first-name does not induce.

4.

Experimental procedure

4.1.

Ethics statement

The study was approved by the local ethical committee (Centre Leon Bérard, Lyon), and subjects gave written informed consent according to the Declaration of Helsinki.

4.2. 3.4.

73

Subjects

Limitations of the study

We carefully controlled most parameters of this passive paradigm. However, some potential confounds or limitations remain. One limitation of our experimental design appears to be the absence of quantitative assessment of attentional load by the primary task. This is an underestimated bias, shared by most studies using a passive paradigm with a primary task like “doing nothing”, “watching a film” or “reading a book”,

Thirty-six healthy right-handed volunteers (18 males, mean age 23 years ± 3) without hearing deficit and without medical, neurological, or psychiatric history participated in the study. In order to decrease stimuli disparities between subjects, one further inclusion criterion was a short first-name: a majority of the subjects had a di-syllabic first-name (27 subjects) and few of them had a first-name with 3 or 4 syllables (9 subjects). Subjects were paid for their participation.

74 4.3.

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Stimuli

The auditory stimuli were spectrally rich tones with a main frequency of 800 Hz and two harmonic partials (1600 and 3200 Hz), the subject's own first-name (OWN) and an unfamiliar first-name (OTHER). For each subject, the choice of OTHER met strict criteria. Firstly, OTHER was chosen to sound equally unfamiliar for all subjects i.e. it was not the first-name of a person in the close environment of the subject. This criterion was controlled thanks to the following procedure a) a questionnaire asking the subject to list the first-names of the persons in their close environment (the questionnaire was filled in a couple of weeks before the recording session), b) a questionnaire presenting OTHER among several first-names and asking the subject to indicate which first-names belonged to a person in his/her close environment (this questionnaire was filled in few days before the recording session). Moreover, first-names belonging to famous people were avoided. Secondly, we chose OTHER so that it could be perceived as different from OWN from the beginning of its utterance, while showing matching acoustic properties, more particularly at its onset. In order to obtain such a result, OTHER was chosen with the same first phoneme as OWN, the first syllables of the two names being however different (see Table 2). Thus, 1) OWN and OTHER differed from their onsets, since their first phonemes were uttered differently through the coarticulation phenomenon (Hardcastle and Hewlett, 1999); 2) they showed a similar energy in their onset, illustrated by the fact that the root mean square (RMS) values of the two stimuli in the first 50 ms were found not different (3321 ± 2074 for OWN, 3259 ± 2235 for OTHER, paired t-test not significant). Additionally, OTHER was chosen to present the same number of syllables as OWN. Altogether, this operating mode allowed to obtain an OTHER stimulus that is perceived as different from OWN from its onset, while minimizing the dissimilarities between the envelops of the acoustic signal plots of both stimuli as illustrated in the example of acoustic plots of one pair of first-names in Fig. 6 (see also Fig. 7, right panel).

First-names were digitally recorded by a neutral masculine voice using Adobe Audition 1.5 (Adobe software). After recording, maximum amplitudes of all stimuli were normalized. The mean durations of OWN (581 ms ± 86) and OTHER (598 ms ± 78) were not significantly different. The 36 pairs of first-names (OWN/OTHER) used in the study are displayed in Table 2.

4.4.

Experimental design

The presentation of the four types of auditory stimuli obeyed the rules of a novelty oddball paradigm. Tones lasting 75 ms and 30 ms (including 5 ms rise/fall times) were used respectively as standard (p = 0.82) and deviant (p = 0.14). OWN and OTHER were presented as novel stimuli. The probability of occurrence was 0.02 for each of them. The stimuli were presented in a pseudo-randomized order so that 1) each deviant followed at least two standards, and 2) each novel followed at least ten standards and/or deviants. Stimulus onset asynchrony was set at 650 ms, except for the standard following a novel, which appeared 1260 ms after the novel onset, whatever the duration of the novel.

4.5.

Procedure

The data presented in this study were recorded during the waking session of a sleep study. Subjects arrived in the lab at 7.00 p.m. after they had eaten. During approximately one hour and a half, electrodes were fixed on their head and face. The subjects selected a DVD among a choice of comedy or action movies. Then subjects were installed in an acoustically dampened and electrically shielded room, earphones were inserted in their ears, and their hearing threshold was assessed using standard stimuli. When the recording session started (10.24 p.m. ± 45 min), stimuli were presented binaurally at 50 dB above the subject's hearing level using the Presentation software (Neurobehavioral Systems). Subjects were instructed to watch the movie (silenced with subtitles) and to ignore the auditory stimuli. The recording session lasted 1 h and 6 ± 9 min.

Table 2 – List of the pairs of own first-name (OWN) and other first-name (OTHER) used for the 36 subjects.

4.6.

Subjects

Twenty-one Ag/AgCl scalp electrodes were manually positioned according to the extended International 10–20 System (Fz, Cz, Pz; FP1, F3, FC1, C3, T3, CP1, P3, M1, O1 and their counterparts on the right hemiscalp). This relatively small number of electrodes was both compatible with sleep recordings and with the use of scalp potentials (SP) and scalp current density (SCD) maps. We concentrated the electrodes around the sites expected to show between first-name effects i.e. central and parietal sites. Contact between skin and electrodes was made using EC2 electrode cream Pactronic (Grass Product Group) and electrodes were fixed on the scalp using the paste TENSIVE (Parker Laboratories, Inc.). The reference electrode was placed on the tip of the nose, and the ground electrode on the forehead. The electro-oculogram (EOG) was recorded from 2 electrodes placed on the supraorbital ridge of the left eye and on the infraorbital ridge of the right eye. Muscle activity (EMG) was recorded from 2 electrodes attached to the chin. Electrode impedance was kept below

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18

OWN

OTHER

Subjects

Fabrice Fabien Lucie Amal Thomas Emeric Clara Maud Simon Maelle Estelle Yoann Etienne Thomas Marie Gaëlle Jackie Marine

Florent Félix Lydie Alice Tanguy Elie Claudia Mylène Samy Mylène Eva Yasid Edgar Tanguy Mégane Gwendy Justine Mylène

19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36

OWN

OTHER

Rémi Florian Geoffrey Clémence Hélène Marie Marion Myriam Boris Elodie Julien Camille Julie Alexandre Wissam Waël Jérémie Clément

Rayan Fernand Gérard Clara Eva Mégane Mégane Maëlle Bernard Erika Joseph Colette Jackie Adrien Walter Wissam Joseph Claudio

Electrophysiological recordings

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75

Fig. 6 – Acoustic signal plot and spectrogram of the own first-name (OWN) and of the unfamiliar first-name (OTHER) for subject 7.

Fig. 7 – Left panel: Event-related potentials (ERPs) to the own first-name (OWN, in red) and to the unfamiliar first-name (OTHER, in black) for individual subjects 4 and 11 at Cz (A) before the inter-subject N1 alignment and (B) after the inter-subject N1 alignment. The grand average of ERPs to OWN and OTHER after N1 alignment is shown in (C). Right panel: Acoustic signal plot of OWN and OTHER for subjects 4 and 11 (D).

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5 kΩ. The electrophysiological data (EEG, EOG and EMG) were continuously recorded via a BrainAmp system (Brain Products GmbH, Germany) with an amplification gain of 12 500, a highpass filter of 0.1 Hz and a sampling rate of 1000 Hz.

4.7.

Event-related potential analysis

Event-related potential (ERP) analysis was undertaken using the ELAN Pack software (Aguera et al., 2011) developed at the “Brain Dynamics and Cognition” team of the Lyon Neuroscience Research Center (Lyon, France) and the Matlab programming tool (Mathworks). ERPs elicited by OWN and OTHER were averaged over an epoch of 900 ms including a pre-stimulus period of 100 ms. Trials with overall electrophysiological signal amplitude exceeding 150 μV were automatically excluded from the averaging. After averaging, the baseline was corrected according to the mean value of the signal during the 100 ms prior to the stimulus. A 30 Hz low-pass digital filter was applied (bidirectional Butterworth, 4th order). Contrary to some authors in the novelty literature (e.g. Escera et al., 2003), but like Holeckova et al. (2006), we choose not to subtract the ERPs to standards from those to novels, because of the large acoustic discrepancy between first-names and tones. Our aim being to compare responses to OWN and OTHER, subtracting responses to standards from both would not have induced different results. The components elicited by OWN and OTHER were visually identified in the grand averaged ERPs. In individual tracings, they were measured at the electrode that showed the largest amplitude in the grand average. The latency and the amplitude of the peak, as well as the mean amplitude, were assessed inside a time-window around the latency of the peak in the grand average. Maps showing the distribution of the scalp potentials (SP) in the time-windows of interest were generated using a spherical surface spline interpolation algorithm (Perrin et al., 1989). Radial scalp current density (SCD) maps (in mA/m3) were obtained by computing the second spatial derivatives of the spline functions used in potential map interpolation (Perrin et al., 1987; Perrin et al., 1989). SCD distributions have the property of being reference-free and of showing sharper peaks than those of the potential maps (Pernier et al., 1988), which may facilitate data interpretation in case of multiple overlapping sources. ERPs elicited by standard and deviant tones are out of the scope of this study. They are displayed in Supplementary Fig. S1.

4.8.

Inter-subject sensory response alignment

For each subject, OWN and OTHER evoked N1 sensory responses at very similar latencies. However, the different first-names of the subjects showed variable onsets and attacks, as illustrated by a large variance in the RMS value of the first 50 ms of the stimuli (3321 ± 2074 for OWN, 3259 ± 2235 for OTHER). This large inter-subject variance of stimulus onset induced a variability in N1 latencies between subjects (Näätänen and Picton, 1987). In order to properly synchronize ERP responses we realigned N1s between subjects a posteriori. For each subject, N1s were visually

identified by an expert (DM), then the maxima of the upward (negative) slope of N1s evoked by OWN and by OTHER were detected. The middle point of these two slope maxima was taken as individual N1 fiducial point. Finally, for each subject, the two ERP responses were shifted by the same amount as to put individual N1 fiducial points at the median latency of the 36 subjects' fiducial points (126 ms, see Fig. 7). This alignment procedure resulted in N1 peak latency around 160 ms for all subjects. In practice, ERPS were shifted by 15 ± 13 ms in average. As a consequence, this operating mode did not substantially modified grand averaged ERPs (see Supplementary Fig. S2).

4.9.

Statistical analysis

Statistical analysis was performed using Wilcoxon matched rank sign tests. First, tests performed at each sampling point and at each electrode allowed to assess 1) the time intervals where the amplitude of the response evoked by each stimulus was significantly different from the base line level and 2) the time intervals where the responses to OWN and to OTHER differed. Time intervals were accepted when at least 15 consecutive points showed different values at p < 0.05 (Guthrie and Buchwald, 1991). Then, the measurements of the different components detected in response to OWN and to OTHER were compared. Finally, in each time-interval of interest, mean scalp potentials and scalp current densities of the responses to OWN and to OTHER were compared at each electrode.

Acknowledgments We thank the members of the Brain Dynamics and Cognition Team for their helpful discussions. The project was supported by the grant ANR-07-JCJC-0095.

Appendix A. Supplementary data Supplementary data to this article can be found online at doi:10.1016/j.brainres.2012.01.072.

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