ERP correlates of auditory goal-directed behavior of younger and older adults in a dynamic speech perception task

ERP correlates of auditory goal-directed behavior of younger and older adults in a dynamic speech perception task

Behavioural Brain Research 278 (2015) 435–445 Contents lists available at ScienceDirect Behavioural Brain Research journal homepage: www.elsevier.co...

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Behavioural Brain Research 278 (2015) 435–445

Contents lists available at ScienceDirect

Behavioural Brain Research journal homepage: www.elsevier.com/locate/bbr

Research report

ERP correlates of auditory goal-directed behavior of younger and older adults in a dynamic speech perception task Stephan Getzmann ∗ , Michael Falkenstein, Edmund Wascher Leibniz Research Centre for Working Environment and Human Factors at the Technical University of Dortmund (IfADo), Ardeystraße 67, D-44139 Dortmund, Germany

h i g h l i g h t s • • • • •

We study speech perception of young and old adults in a dynamic multi-talker scenario. We analyzed behavioral performance and event-related brain potentials. Sudden shifts in speaker location decrease performance especially in the old group. This disadvantage comes along with delayed attentional re-orienting. Age-related declines in speech perception are partly based on cognitive slowing.

a r t i c l e

i n f o

Article history: Received 12 August 2014 Received in revised form 15 October 2014 Accepted 20 October 2014 Available online 29 October 2014 Keywords: Aging Speech perception “Cocktail Party” scenario Mismatch negativity (MMN) P3a Reorienting negativity (RON)

a b s t r a c t The ability to understand speech under adverse listening conditions deteriorates with age. In addition to genuine hearing deficits, age-related declines in attentional and inhibitory control are assumed to contribute to these difficulties. Here, the impact of task-irrelevant distractors on speech perception was studied in 28 younger and 24 older participants in a simulated “cocktail party” scenario. In a twoalternative forced-choice word discrimination task, the participants responded to a rapid succession of short speech stimuli (“on” and “off”) that was presented at a frequent standard location or at a rare deviant location in silence or with a concurrent distractor speaker. Behavioral responses and eventrelated potentials (mismatch negativity MMN, P3a, and reorienting negativity RON) were analyzed to study the interplay of distraction, orientation, and refocusing in the presence of changes in target location. While shifts in target location decreased performance of both age groups, this effect was more pronounced in the older group. Especially in the distractor condition, the electrophysiological measures indicated a delayed attention capture and a delayed re-focussing of attention toward the task-relevant stimulus feature in the older group, relative to the young group. In sum, the results suggest that a delay in the attention switching mechanism contribute to the age-related difficulties in speech perception in dynamic listening situations with multiple speakers. © 2014 Elsevier B.V. All rights reserved.

1. Introduction Speech comprehension under “cocktail party” conditions [14] poses a great challenge to the human auditory system: When confronted with more than one speaker at once, it is necessary to perceptually segregate the speech information of interest from background noise and to focus auditory attention on the target speaker, while inhibiting the information from concurrent speakers (for review, see [19]). Listening under these conditions is especially

∗ Corresponding author. Tel.: +49 231 1084 338; fax: +49 231 1084 401. E-mail address: [email protected] (S. Getzmann). http://dx.doi.org/10.1016/j.bbr.2014.10.026 0166-4328/© 2014 Elsevier B.V. All rights reserved.

difficult when the spatial distance between target and concurrent speakers is small (e.g., [4,11]). The situation should be even more challenging when target and concurrent speakers are not static, but dynamically change their spatial positions like in a normal conversation of two or more speakers. Here, it is important to quickly switch between changing speakers and to flexibly adapt auditory attention to the current speaker of interest. It is known that speech comprehension under “cocktail party” conditions becomes more difficult in aging. This is mainly due to changes in cochlear, retrocochlear, and central auditory processing causing an overall loss of auditory information (for reviews, see [25,41]). In addition, age-related declines at a higher cognitive level, such as working memory capacity and information processing

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speed, reduces the ability to understand language in multi-talker situations (for review, see [12]). In particular, according to the inhibitory deficit hypothesis [35], attentional and inhibitory control usually decline, so that older adults are more readily distracted by task-irrelevant stimuli than younger adults (for review, see [45]). Thus, in a dynamic “cocktail party” situation, deficits in attentional control may hinder the fast adaptation of auditory attention to a current target speaker, making older adults more susceptible to a distraction by sudden changes in speaker constellations. The aim of the present study was to address the role of distraction by unexpected changes in the auditory environment and consequences of age-related deficits in attentional control for listening in dynamic “cocktail party” situations. A speechcomprehension task was hereto employed. It was based on a well-characterized auditory distraction paradigm used to investigate the cognitive sub-processes underlying attentional and inhibitory control in a number of previous studies [66,21,65,60]. In this “classical” paradigm, a sequence of repeated (standard) tones is intermixed with occasional irregular (deviant) tones violating the repetition. When subjects have to respond to a relevant tone features (e.g., the tone duration), the unexpected occurrence of a deviant (but task-irrelevant) tone feature (e.g., a variation in tone pitch or location) usually decreases performance, indicated by a decrease in correct responses and an increase in reaction times (RTs). This effect has been interpreted within the theoretical framework of a three-stage model of distraction, in which the irregular deviant feature triggers a deviance detection–attention switching–attention re-focussing cycle (e.g., [66,21]): In the first stage of regularity extraction and deviance-detection the taskrelevant information is filtered out of the stream of on-going stimulation, and deviant and novel information is automatically detected in terms of violations of regularities in sensory memory buffers. In the second stage of attention-switching the deviant information leads to an involuntary shift of attention which is then – in a third stage of re-orientation and re-focussing – compensated for by mechanisms restoring the attention-set relevant for a given task. Neurophysiological evidence of these cognitive sub-processes are given by event-related potentials (ERPs): While both standard and deviant stimuli usually produce a fronto-central N1–P2–N2 complex, reflecting automatic processing of the sensory input (N1; [51]), allocation of attention and stimulus evaluation (P2; [52,58]), and executive cognitive control functions (N2b; for review, see [26]), the difference ERPs (deviant minus standard) typically reveal a chain of deviant-related potentials, consisting of the mismatch negativity (MMN), P3a, and reorienting negativity (RON) (e.g., [40]). The MMN ([49]; for review, see [50]) is a correlate of pre-attentive deviance detection [48,68], the fronto-central P3a [27] reflects an involuntary attention-switching mechanism [21,27,43,64], and the RON [65,66] is assumed to indicate re-allocation of attention to the relevant task after distraction by a deviant features (for empirical evidence: [6,38]). As could be expected from the inhibitory deficit hypothesis, older adults are more distracted by irregular deviant stimuli than younger adults [46,15,39,61]. The analysis of the deviant-related ERPs revealed that different cognitive subprocesses contribute to this increase in distractibility, comprising deficits in encoding or retention of sensory information [1,9,15,17,42,54,61] and in attentional orienting [18,28,46,39]. Also, a delayed and incomplete re-orientation of attention to the relevant stimulus feature has been observed [46,39,31,32]. In order to adapt the theoretical framework of the distraction–orientation–refocusing cycle to a dynamic “cocktail party” situation, a word discrimination task was employed here. A rapid sequence of monosyllabic words was presented from either a frequent (standard) location or a rare (deviant) location. The participants had to respond to the words as fast

as possible, while ignoring the irregular shifts of the speaker’s position. Thus, they had to attend to the task-relevant feature (i.e., the word content), while ignoring the task-irrelevant feature (i.e., the speaker position). The performance of younger and older adults was measured in a single-speech baseline condition, in which only the target speaker was present, and in two multispeech conditions: Here, concurrent speech information was presented simultaneously to the target speech by a second speaker located either far away or near the target speaker. It was expected that older participants suffer more from irregular shifts of the target speaker than the younger ones, and that the decrease in performance was especially pronounced when the concurrent speaker was close to the target speaker. To study the sources of performance differences, the ERPs to speech stimuli at standard and deviant positions were contrasted and analyzed in both age groups, and for the single-speech and multi-speech conditions. 2. Methods 2.1. Participants A total of 52 volunteers took part in the study, consisting of 28 younger (15 female; mean age 25.0 years; range 19–32 years) and 24 middle-aged and older adults (12 female; mean age 62.6 years; range 55–72 years). The young participants were recruited from local colleges, while the older participants were recruited through newspaper advertisements. All participants reported to be righthanded and healthy, free of medication during the experimental sessions, and without any history of neurological, psychiatric, or chronic somatic problems. The hearing thresholds of the participants were examined by pure-tone audiometry (Oscilla USB 330; Inmedico, Lystrup, Denmark) at 125–8000 Hz. Except mild to moderate presbyacusis in the older group, the audiograms of all but one participants were within a defined tolerance zone, indicating normal hearing below 4000 Hz (thresholds better than 30 dB hearing loss). One participant (male; 72 years old) was excluded from the analysis due to a pronounced hearing loss (49 dB). All participants gave their written informed consent before any study protocol was commenced. The study conformed to the Code of Ethics of the World Medical Association (Declaration of Helsinki) and was approved by the local Ethical Committee of the Leibniz Research Centre for Working Environment and Human Factors, Dortmund, Germany. 2.2. Stimuli, task, and procedure The experiment took place in a dimly lit, electrically shielded, and sound-proof room (5.0 × 3.3 × 2.4 m3 ) with pyramid-shaped foam acoustic panel on ceiling and walls, and a sound-absorbing woolen carpet on the floor. The ambient background noise level was below 20 dB(A) SPL. During the experiment, the subject was seated on a vertically adjustable armchair. The position of the head was held constant by a chin rest. The target stimuli consisted of monosyllabic German words (“an” [“on”] and “ab” [“off”]), the concurrent stimuli were spoken numerals (“eins” [one] to “zehn” [ten]). The words were spoken by monolingual native German speakers without any dialect or speech disorders, the target stimuli by a female, the concurrent stimuli by a male (voice fundamental frequencies 170 Hz and 119 Hz, respectively). The stimuli were digitally recorded in a sound-proof and anechoic chamber, using a freestanding microphone (Beyerdynamic MCE 91) and a mixing console (Mackie 1202-VLZpro, sampling rate 48 kHz). The sound stimuli were processed offline using CoolEdit 2000 (Syntrillium Software Co., Phoenix, AZ, USA). The duration of all speech stimuli was adjusted to 300 ms. To make sure that younger and older

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(deviant at 75◦ ), at 75◦ (deviant 15◦ ), at −15◦ (deviant at −75◦ ), or at −75◦ (deviant at −15◦ ). Thus, both standard and deviant stimuli were located within the left or right hemispace, and the deviant stimulus was horizontally shifted by 60◦ to the left or the right of the standard position. The spatial arrangements of standard and deviant locations was kept constant for each participant, but balanced across participants. The sequence of standards and deviants was pseudo-randomized in a way that each deviant location was followed by at least three standard locations. In the no-distractor condition, the sequence of target words was presented in silence. In the multi-speech conditions, the target words were presented simultaneously with the concurrent words. The numerals were presented in a pseudo-randomized order. In the far-distractor condition, the speaker of the concurrent words was located in the hemispace opposite to the target speaker: at −45◦ when the target speaker was at 15◦ and 75◦ , and at 45◦ when the target speaker was at −15◦ and −75◦ . In the near-distractor condition, the speaker of the concurrent words was located in the same hemispace as the target speaker, midway between the standard and deviant locations (i.e., at 45◦ when the target speaker was located at 15◦ and 75◦ , and at −45◦ when the target speaker was located at −15◦ and −75◦ ). Before the test started, the participants carried out a short training until the task was familiar. It was also tested whether all participants were able to easily understand the words, and to perceive the shifts in spatial position. Then, the participants completed 480 trials for each distractor condition, consisting of 400 stimuli at standard locations (200 “an” and 200 “ab”) and 80 stimuli at deviant locations (40 “an” and 40 “ab”). The distractor conditions were presented blockwise, and counterbalanced across the participants. The timing of the stimuli and the recording of the participants’ responses were controlled by custom-written software. 2.3. EEG recording and pre-processing

Fig. 1. (A) Experimental set-up with six loudspeaker in the participant’s left and right hemispace, constituting standard and deviant positions (±75◦ and ±15◦ ) and distractor positions (±45◦ ). (B) Target speech stimuli “an” and “ab” were presented at frequent standard locations and rare deviant locations (here: −15◦ and −75◦ ). The target speech was presented in isolation (no distractor), or in the presence of concurrent speech stimuli (“1” to “9”) from a far or near position (here: far distractor at 45◦ or near distractor at −45◦ ), relative to the target speech position. Standard stimuli (solid) and deviant stimuli (dotted) are encircled.

participants could easily perceive the speech stimuli, the sound level was well above the hearing threshold at about 65 dB(A). The stimuli were converted to analogue form via a computer-controlled external soundcard (Sound Blaster Audigy 2 NX, Creative Labs, Singapore) and presented by six broad-band loudspeakers (SC 5.9; Visaton, Haan, Germany) that were mounted in front of the subject at a distance of 1.5 m from the center of the head (Fig. 1A). The two target stimuli (“an” and “ab”) were presented equiprobably and with a constant inter-stimulus interval of 1100 ms, following a pseudo-randomized order (Fig. 1B). In a two-alternative forced-choice word discrimination task, the participants were instructed to press the upper response button of a response box for the word “an” (i.e., “switching on”), and the lower button for the word “ab” (i.e., “switching off”), using the index and the middle finger of the dominant hand. The response box was held in the participant’s hands. They had to respond in a fast but accurate manner. The participants were instructed to keep their eyes open and to focus on a visual fixation point in front of them. No feedback was given at any time during the experiment. The speech stimuli were presented either from a frequent standard location (83.33%) or a rare deviant location (16.66%). The standards were located at 15◦

The continuous EEG was sampled at 2048 Hz using 64 electrodes and a BioSemi amplifier (Active Two; Biosemi, Amsterdam, Netherlands). Electrode positions were based on the International 10–10 system. The amplifier bandpass was 0.01–140 Hz. Horizontal and vertical eye positions were recorded by electrooculography (EOG) using six electrodes positioned around both eyes. Two additional electrodes were placed on the left and right mastoids. Electrode impedance was kept below 10 k. The raw data were downscaled offline to a sampling rate of 1000 Hz, digitally band-pass filtered (cut-off frequencies 0.5 and 25 Hz; slopes 48 dB/octave), and re-referenced to the average of the mastoid electrodes. The data were segmented into 1350-ms stimulus-locked epochs covering the period from −100 to 1250 ms relative to stimulus onset. Data were then corrected for ocular artifacts using the Gratton, Coles, and Donchin procedure [33]. Individual epochs exceeding a maximum–minimum difference of 200 ␮V and a maximum voltage step of 50 ␮V per sampling point were excluded from further analysis (automatic artifact rejection as implemented in the BrainVision Analyzer software, Version 1.05; Brain Products, Gilching, Germany). The remaining epochs were baseline corrected with reference to a 100-ms prestimulus window, and averaged for each participant, separately for epochs with deviant and standard locations. Trials were averaged across the two target words (“an” and “ab”), and across numerals in the multi-speech conditions. Finally, difference waves were calculated (deviant minus standard) to analyze the deviance-related MMN, P3a, and RON components. 2.4. Data analysis Behavioral and ERP data were analyzed for standard and deviant stimuli, and for the single-speech and multi-speech conditions. In

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order to minimize potential effects of post-deviance distraction (in which the processing of standard stimuli is affected by a preceding deviant stimulus; e.g., [32]) only standard trials preceding a deviant were analyzed. RT was defined as the time between the onset of the target stimulus and the push of a response button. Given that the “an” and “ab” stimuli did not differ until 100 ms after stimulus onset, RTs were corrected for this time lag. Individual RTs of less than 100 ms and more than 1200 ms, as well as error trials were excluded from further analysis. Rates of correct responses and mean RTs were subjected to three-way analyses of variance (ANOVAs) with between-subject factor age (younger, older) and within-subject factors position (deviant, standard) and distractor (no, far, near). In addition, the relative changes in RTs and in correct responses were computed and submitted to two-way ANOVAs with between-subject factor age and within-subject factor distractor. The ERP analysis was restricted to midline electrodes (Fz, FCz, Cz, and Pz) chosen to be commensurate with previous knowledge of the topographical scalp distribution of specific ERPs (review: [5,51,56]), indicating that the P1, P2, MMN, N2b, P3a, and RON typically peak over fronto-central areas (FCz), the N1 over central areas (Cz), and the P3b over parietal areas (Pz) of the scalp. Peak amplitudes and latencies were defined as their local maximum positivity or negativity within a particular latency window for standard and deviant trials (P1 at FCz: 20–120 ms; N1 at Cz: 75–175 ms; P2 at FCz: 150–250 ms; P3b at Pz: 400–800 ms; after speech onset) and for the difference waves (MMN: 100–200 ms; P3a: 200–400 ms; RON: 300–700 ms; all at FCz). In addition, the amplitude of the N2b (visible only in the young group, see below) was determined for each participant as the mean amplitude of a 40-ms period centered at the average peak latency of the young group (290 ms at FCz). The amplitudes and peak latencies of P1, N1, P2, and P3b as well as the N2b amplitudes were subjected to three-way ANOVAs with between-subject factor age and within-subject factors position and distractor. The amplitudes and peak latencies of the deviance-related MMN, P3a, and RON were subjected to two-way ANOVAs with between-subject factor age and within-subject factor distractor. Levene’s test was used to assess the homogeneity of variance, and the degrees of freedom were adjusted if variances were unequal. Effect sizes were computed to provide a more accurate interpretation of the practical significance of the findings, using the partial eta-squared (p 2 ) coefficient.

3. Results 3.1. Behavioral data 3.1.1. Response accuracy The younger participants made a larger number of correct responses than the older ones (93.4% vs. 90.3%), while the main effect of age slightly failed to reach significance (F1, 49 = 3.97; p = 0.052; p 2 = 0.08). Also, there were more correct responses without distractor (93.8%) than with the far distractor (91.1%) and the near distractor (90.6%) (main effect distractor: F2, 98 = 6.85; p < 0.01; p 2 = 0.12). Post-hoc t-tests indicated significant differences between the no-distractor and far-distractor conditions, and between the no-distractor and near-distractor conditions (both p < 0.05), while far-distractor and near-distractor conditions did not differ (p > 0.05). A deviant target location did not decrease the overall rate of correct responses (no main effect position: F1, 49 = 0.44; p > 0.05; p 2 = 0.01). Also, there were no interactions of age and distractor (F2, 98 = 2.85) or age and position (F1, 49 = 2.27), or distractor and position (F2, 98 = 0.55; all p > 0.05; all p 2 < 0.06). However, a significant interaction of age, position, and distractor occurred (F2, 98 = 3.49; p < 0.05; p 2 = 0.07), suggesting that the effect of age was most pronounced when the target appeared at the deviant

location, and when the distractor stimulus was close to the target location (Fig. 2A). In order to further analyze this interaction, the change in the rate of correct responses at the deviant location, relative to the standard location, was computed and submitted to a two-way ANOVA with between-subject factor age and within-subject factor distractor. The older participants did not differ from the younger ones in their overall change in rate of correct responses (no main effect age: F1, 49 = 2.30; p > 0.05; p 2 = 0.05). In fact, the performance of both age groups was less affected when there was no distractor and when the distractor was far away (Fig. 2C). However, when the distractor was near, the older group showed a decrease in performance (−3.1%), whereas the younger group showed a slight increase (1.6%) (interaction age and distractor: F2, 98 = 3.27; p < 0.05; p 2 = 0.06). This difference in the change of accuracy was significant (t49 = 2.17; p < 0.05). 3.1.2. Response times The older participants responded significantly later than the younger ones (482 ms vs. 424 ms; main effect age: F1, 49 = 22.48; p < 0.01; p 2 = 0.32). Both age groups responded later when the distractor was present (far distractor: 456 ms; near distractor: 463 ms) than without distractor (440 ms) (main effect distractor: F2, 98 = 29.63; p < 0.001; p 2 = 0.38). Post-hoc t-tests indicated significant differences between all distractor conditions (all p < 0.05). Both age groups also responded later to target words at deviant locations, relative to standard locations (462 ms vs. 444 ms; main effect position: F1, 49 = 54.23; p < 0.001; p 2 = 0.53). This effect was stronger in the older group (+25 ms) than in the younger group (+11 ms) (interaction age and position: F1, 49 = 7.54; p < 0.01; p 2 = 0.13; Fig. 2B). There were no interactions of age and distractor (F2, 98 = 0.70) or age, distractor, and position (F2, 98 = 0.44; both p > 0.05; both p 2 < 0.02). Accordingly, the analysis of the relative change in RTs at deviant relative to standard locations did not indicate an interaction of age and distractor (F2, 98 = 0.28; p > 0.05; p 2 = 0.01). However, there was a larger overall increase in RTs in the older group than in the younger group (4.4% vs. 2.2%; main effect age: F1, 49 = 5.84; p < 0.05; p 2 = 0.11; Fig. 2D). In sum, the older participants made slightly more errors and responded later than the younger ones. A deviant target location caused a decrease in performance, indicated by an increase in RTs. This increase in RTs was stronger in the older group than in the younger group. Also, a concurrent speech stimulus decreased performance, indicated by an increase in RTs and a decrease in the rate of correct responses. When the concurrent speech stimulus was close to the target speech location, the rate of correct responses at deviant locations decreased in the older group, but remained stable in the younger group. 3.2. ERP of standard and deviant locations The speech stimuli produced a typical fronto-central P1-N1-P2 complex and a parietal P3b, peaking at 61 ms, 119 ms, 205 ms, and 569 ms, respectively (averaged across age groups and conditions; Fig. 3, left and central panel). The young group showed an additional distinct negative fronto-central peak (N2b) at about 290 ms that was absent in the older group. In contrast, the older group showed a pronounced late frontal positivity. In the following sections, significant effects of age on these ERP components and significant interactions of age and position or distractor are described. 3.2.1. P1 The older group showed an overall larger P1 amplitude than the younger ones (3.6 ␮V vs. 2.5 ␮V; main effect of age: F1, 49 = 6.84; p < 0.05; p 2 = 0.12), but there were no interactions of age and

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Fig. 2. Rates of correct responses (A) and reaction times (B) for younger and older participants, shown for standard and deviant locations and for no-distractor, far-distractor, and near-distractor conditions. Change in rates of correct responses (C) and in reaction times (D) at deviant locations, relative to standard locations. Error bars indicate standard errors across participants (younger: N = 28; older: N = 23).

position or distractor. Also, there was no effect of age on P1 latency (all p > 0.05; all p 2 < 0.03). 3.2.2. N1 The N1 latency was slightly delayed in the older group (122 ms vs. 117 ms; main effect of age: F1, 49 = 5.04; p < 0.05; p 2 = 0.09). There were no main effects of age on N1 amplitude, and no interactions of age and position or distractor on N1 amplitude or latency (all p > 0.05; all p 2 < 0.05). 3.2.3. P2 The P2 latency was delayed in the older group (214 ms vs. 197 ms; main effect of age: F1, 49 = 22.33; p < 0.001; p 2 = 0.32). The P2 amplitude appeared to be slightly larger in the older group (4.8 ␮V vs. 3.1 ␮V; main effect of age: F1, 49 = 3.27; p = 0.08; p 2 = 0.06), but this effect was modulated by an age × distractor interaction (F2, 98 = 3.80; p < 0.05; p 2 = 0.07): The younger group showed a clear reduction in P2 amplitude when distractor was present, while the older group showed a marginal reduction (cf. Fig. 3). The P2 reduction of the younger group occurred mainly at the deviant locations, while the P2 remained constant at the target locations irrespective of distractor presence (interaction of distractor and position: F2, 98 = 27.71; p < 0.001; p 2 = 0.36). The differences between age groups were confirmed by additional ANOVAs separately for the younger and older groups. In the younger group, a main effect of distractor occurred (F2, 54 = 17.84; p < 0.001; p 2 = 0.40), and post-hoc t-tests indicated significant differences in the P2 amplitude between the far-distractor and

near-distractor conditions (both 2.4 ␮V) and the no-distractor condition (4.4 ␮V; both p < 0.001). In the older group, the interaction did not reach significance (F2, 44 = 2.51; p < 0.05; p 2 = 0.10). There were no further interactions (all p > 0.05; all p 2 < 0.02). 3.2.4. N2b Only the young group showed a pronounced N2b component, while the older group showed a slight negative deflection relative to the P2 positivity (−3.7 ␮V vs. 3.6 ␮V; cf. Fig. 3; main effect of age: F1, 49 = 62.67; p < 0.001; p 2 = 0.56). There were no interactions (all p > 0.05; all p 2 < 0.02). 3.2.5. P3b The P3b was delayed in the older group (589 ms vs. 549 ms: main effect of age: F1, 49 = 4.63; p < 0.05; p 2 = 0.09). There were no main effects of age on P3b amplitude and no interactions (all p > 0.05; all p 2 < 0.04). 3.2.6. Late positivity In order to analyze the increased frontal late positivity (LP) observed in the older group, the mean amplitudes at electrode positions Fz, FCz, Cz, and Pz were computed in the time range between 300 and 600 ms and submitted to a four-way ANOVA with betweensubject factor age and within-subject factors position, distractor, and frontality (Fz, FCz, Cz, Pz). The older group showed much stronger amplitudes than the younger ones (3.4 ␮V vs. −0.1 ␮V; main effect of age: F1, 49 = 22.57; p < 0.001; p 2 = 0.32). Moreover, the two groups differed especially over frontal areas, where the

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Fig. 3. Grand average ERPs of target words at frequent standard locations (left) and at rare deviant locations (central), and difference waves (deviant minus standard locations; right), shown for no-distractor, far-distractor, and near-distractor conditions, and younger and older participants at fronto-central ((A) FCz) and parietal ((B) Pz) electrode positions. ERP components (P1, N1, P2, N2b, P3a, P3b, late positivity LP MMN, and RON) are marked.

older participants showed a positivity and the younger ones a negativity. No differences occurred over parietal areas (Fig. 4; interaction of age and frontality: F3, 147 = 52.21; p < 0.001; p 2 = 0.52). There were no further interactions (all p > 0.05; all p 2 < 0.04).

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Electrode Fig. 4. Mean amplitudes of younger and older participants at frontal (Fz), frontocentral (FCz), central (Cz), and parietal (Pz) electrode positions in the time range 300–600 ms. Error bars are standard errors across participants (younger: N = 28; older: N = 23).

3.3. Deviance-related ERPs The difference waves showed a fronto-central MMN, P3a, and RON complex, peaking at 157 ms, 293 ms, and 496 ms, respectively (averaged across age groups and conditions; Fig. 3, right panel). In the following sections, significant effects of age on these deviancerelated ERPs are reported. 3.3.1. MMN There were no main effects of age and no interactions of age and distractor on MMN amplitude or latency (all p > 0.05; all p 2 < 0.05). Irrespective of age, the MMN latency was larger in the neardistractor condition (180 ms) than in the far-distractor (151 ms) and no-distractor (139 ms) conditions (Fig. 5A; main effect of condition: F2, 98 = 26.48; p < 0.001; p 2 = 0.35). Post-hoc t-tests indicated significant latency differences between near-distractor and fardistractor conditions, and between near-distractor condition and no-distractor conditions (both p < 0.001), while the far-distractor and no-distractor condition did not differ. 3.3.2. P3a The P3a was delayed in the older group (307 ms vs. 277 ms; main effect of age: F1, 49 = 9.19; p < 0.01; p 2 = 0.16). Also, the P3a was delayed in the near-distractor condition (338 ms) compared to the far-distractor condition (291 ms) and the no-distractor

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Fig. 5. Mean amplitudes and peak latencies of the MMN (A), P3a (B), and RON (C), shown for no-distractor, far-distractor, and near-distractor conditions, and younger and older participants. Error bars are standard errors across participants (younger: N = 28; older: N = 23).

condition (250 ms) (main effect of condition: F2, 98 = 65.78; p < 0.001; p 2 = 0.57). Post-hoc t-tests indicated significant latency differences between all three distractor conditions (all p < 0.001). Moreover, there was a significant age × distractor interaction (F2, 98 = 4.65; p < 0.05; p 2 = 0.09), suggesting that the increase in P3a latency in the near-distractor condition was more pronounced in older, than younger, participants (Fig. 5B). In fact, additional post-hoc t-tests indicated significant latency differences between the two age groups only in the near-distractor condition (t49 = 4.09; p < 0.001), but not in the far-distractor condition (t49 = 1.26; p > 0.05) or in the no-distractor condition (t49 = 1.11; p > 0.05). The P3a amplitude was overall reduced in the older group (3.8 ␮V vs. 4.9 ␮V; main effect of age: F1, 49 = 4.52; p < 0.05;

p 2 = 0.08). Moreover, it was higher in the no-distractor condition (5.5 ␮V) than in the far-distractor and near-distractor (both 3.8 ␮V) conditions (Fig. 5B; main effect of condition: F2, 98 = 13.43; p < 0.001; p 2 = 0.22). However, there was no age × distractor interaction on P3a amplitudes (F2, 98 = 1.25; p > 0.05; p 2 = 0.03). 3.3.3. RON The RON was delayed in the older group (540 ms vs. 449 ms; main effect of age: F1, 49 = 40.63; p < 0.001; p 2 = 0.45). Also, the RON was delayed in the near-distractor condition (520 ms) compared to the far-distractor condition (486 ms) and the no-distractor (477 ms) condition (Fig. 5C; main effect of condition: F2, 98 = 7.71; p < 0.01; p 2 = 0.14). Post-hoc t-tests indicated significant latency differences between the near-distractor condition and the two

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RON latency [ms] Fig. 6. Pearson correlations of changes in inverse efficiency (the error rate adjusted reaction times at deviant locations relative to standard locations) against RON latencies in the near-distractor condition for each participant, shown separately for older (N = 23) and younger (N = 28) participants with linear regression lines.

other distractor conditions (p < 0.05), while the far-distractor and no-distractor conditions did not differ. There was also no age × distractor interaction on RON latency (F2, 98 = 0.02; p > 0.05; p 2 = 0.01). There were no main effects of age on RON amplitudes and no interactions (all p > 0.05; all p 2 < 0.02). 3.4. Correlational analyses In order to test whether the decrease in performance found in the older group at deviant locations with the near distractor was related to the observed increase in P3a and RON latencies in this age group, correlations between changes in performance in the near-distractor condition and the latencies of P3a and RON were computed, separately for the younger and older participants. To this aim (and to reduce the number of correlation calculations) a combined measure of RT and accuracy was employed—the inverse efficiency (IE). The IE score (expressed in ms) was calculated by dividing the RT by the rate of correct responses, thus accounting for a possible tradeoff between speed and accuracy [69]. There was a significant correlation of RON latency and change in IE in the older group (r = 0.52; p = 0.011; two-tailed), but not in the younger group (r = −0.15; p > 0.05). The correlation coefficients of the two age groups differed significantly (Fisher’s Z = 2.42, two-tailed p < 0.05). Thus, a stronger decrease in performance of older participants came along with increased RON latencies, while performance of younger participants was not related to RON latency (Fig. 6). The correlation of P3a latency and change in IE did not reach statistical significance in the older group (r = 0.34) or in the younger group (r = −0.23; both p > 0.05). 4. Discussion Aim of the present study was to study the impact of irregular shifts of a target speaker location on speech perception of younger and older adults in a simulated dynamic “cocktail party” scenario. In general, irregular shifts of the target speaker location caused an increase in RTs that was stronger in the older than younger group. In addition, the presence of a concurrent second speaker increased RTs, irrespective of whether the second speaker was far away or near the target speaker locations. A shift of the target speaker location did not change the response accuracy, at least as long as there was no concurrent speaker or as long as the concurrent speaker was far away from the target locations. However, when the concurrent speaker was close to the target speech location, the older participants showed a decrease in the rate of correct responses, while the performance of the younger participants remained stable. Thus, it

appeared that older participants suffer more from irregular shifts in the relevant speaker location when concurrent (task-irrelevant) speech information is close to the target speaker. These findings are in good accordance with common reports of older adults having less problems in speech comprehension in a quiet environment with only one speaker talking at once. In noisy environments, however, they run into difficulties committing more errors when a sudden change in speaker location occurred. The present results also correspond to the findings of previous auditory distraction studies that found a higher degree of distraction in older adults when an irregular change in a task-irrelevant feature of a target stimulus occurred (e.g., [46,15,39,61]). The analysis of the ERPs revealed neurophysiological correlates of this decline in performance. There was an overall increase in N1, P2, and P3b latencies of the older group that was not modulated by target location or distractor condition. Together with the generally prolonged RTs in the older group, the increase in ERP latencies is in line with the previous findings (e.g., [3]) and the general cognitive slowing hypothesis [62,63]. Interestingly, the older group showed a nearly linear increase in peak latencies relative to the younger group that started from +5 ms (at the time of N1) to +17 ms (P2) and +40 ms (P3b) up to +61 ms (in RTs). The analysis of the deviance-related ERPs revealed even more important results: While there were no latency differences in MMN, the P3a was delayed in the older group when the distractor was close to the target speech location. The delay in P3a latency proceeded to the RON latency that was delayed in the older group in all distractor conditions. This pattern of results is in line with findings of a recent study where auditory distraction was studied in a more abstract experimental setting, and where older participants mainly showed an increase in P3a and RON latencies, rather than in MMN latency [31]. Within the theoretical framework of the three-stage model of distraction (e.g., [66,21]), the MMN is regarded as a correlate of automatic detection of changes in the acoustic environment that may induce a switch of attention to potentially relevant information [48,50,68]. The present results did not provide firm evidence for an effect of age on these early stages of regularity extraction and deviance detection. In fact, the processing of the shift in spatial position on an early, rather pre-attentive level appeared to be quite similar in younger and older participants. However, the switch of attention toward the deviant spatial information (reflected by the P3a; [21,27,43,64]) appeared to be delayed in the older group in the critical condition. There is evidence that the P3a does not reflect a single process of attention switching, but (at least) consists of two subcomponents that mirror two different aspects of attentional control in the context of distraction: An early subcomponent reflecting an automatic disengagement of attention away from the present task-set, and a later subcomponent reflecting controlled attention (e.g., [22,7], 2014). The present results revealed a bi-phasic P3a with two peaks around 300 ms especially in the older group while the younger group showed a sole P3a peak (cf. Fig. 3). Assuming two distinct P3a subprocesses, increased latencies in older group could have decomposed these subprocesses that coincided in the younger group. Decomposition of distinct subprocesses could also be the reason for the overall reduced P3a amplitude in the older group. Moreover, a closer inspection of the P3a peaks of the two age groups (especially in the no-distractor condition) suggests the later subprocess of controlled attention to be more delayed relative to the younger group than the earlier subprocess of automatic disengagement of attention. Alternatively, in the near-distractor condition the prolongation of the P3a in the older group could be due to a reduced early phase of automatic disengagement of attention. These conclusions, however, appear quite speculative and require further examination.

S. Getzmann et al. / Behavioural Brain Research 278 (2015) 435–445

Finally, the re-focussing of attention toward the task-relevant stimulus feature (indicated by the RON; [65,66,6]) was delayed in the older group, suggesting that the attention-switching mechanism took longer and that elderly required more time to overcome distraction than younger participants [8]. Specifically, the switch of attention away from the task-irrelevant change in spatial position back to the task-relevant word content took obviously more time in the older group. As a consequence, especially in the critical condition, the older participants performed worse than the younger ones. In line with this notion, there was a correlation of RON latency and decline in performance, suggesting that a later RON was related to a stronger distraction effect in the older group. No such correlation was found in the younger group, suggesting that their performance was less associated with the period of re-orientation. The notion that age differentially affected MMN, P3a, and RON latencies corroborates previous findings indicating that the three stage of the deviance detection–attention switching–attention refocussing cycle are not strongly coupled [59,40]. On the other hand, the peak latencies of all these ERPs were delayed when the distractor was near. Thus, the distractor obviously slowed down each stage of the deviance detection–attention switching–attention refocussing cycle, resulting in a delay in speech comprehension in both younger and older adults. The older ones, however, needed more time to recover from the distraction. Taken together, these findings suggest that difficulties in speech comprehension that older adults encounter in dynamic “cocktail party” situations could be partly based on timing problems in attentional control. Besides peak latencies, the distractor also affected the P3a amplitude, together with the related P2 amplitude in deviant trials. These amplitudes were smaller when the distractor was present. The P2 has been related to processes of stimulus evaluation [58] and attention allocation [52,71]. The deviance-related P3a is assumed to be a correlate of an attention-switching mechanism [21,27,43,64,57], with larger amplitudes associated with more orientation toward the deviant event. On a phenomenological level, it appears plausible that the shift of target location was less salient when accompanied by a concurrent sound. On a cognitive level, however, the decrease in P2/P3a amplitudes could be interpreted in a way that the distractor suppressed attention allocation toward the shift of target location. In addition to differences in the deviant-related ERPs, the analysis of the ERPs in standard and deviant trials indicated age-related characteristics in speech processing. Firstly, there was a stronger P1 amplitude in the older group. The P1 is elicited very early in the auditory perceptual processing stream [34], and is mainly triggered by the physical characteristics of a stimulus. However, there is also evidence that the P1 is a correlate of early sensory selection (e.g., [36]). Specifically, within the sensory “gain control” model of [37], the P1 is assumed to be an early ERP index of attentional control that interacts with bottom-up sensory processing. A larger P1 amplitude in the older adults has been found in a number of previous studies (e.g., [24]), and could indicate a modality-unspecific augmentation of early stimulus evaluation mechanisms ([67]; for review, see [44]). In the present context of speech perception, the strong P1 component at least indicates that the early process of stimulus intake is unimpaired in the older participants. At a later point in time the differences in ERPs became even more evident: While the younger group showed a pronounced N2b component, the older group showed an ongoing fronto-central positivity (cf. Fig. 3). The N2b is assumed to index processes of cognitive control (for reviews, see [26,53]) and has been related to conflict processing or inhibitory control of irrelevant information (e.g., [47,23,10]). Age-related differences in N2b have been already found in previous studies [3,55,16,2,10,70,32]. In particular, increases in N2b latency and decreases in N2b amplitude have been interpreted as a weakening of controlled processes requiring selective attention

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with increasing age. Specifically, the lack of N2b in elderly has been interpreted as a decrease in the capability to suppress successfully the processing of irrelevant information [10], in line with the inhibitory deficit hypothesis [35]. In the present context, the strong N2b in the young group could reflect the deep processing of the target speech and the inhibition of the concurrent speech stimuli. Interestingly, the age-related decline in N2b was observed in both standard and deviant trials, and irrespective of whether the distractor was present or not. Thus, the N2b decline was not specifically related to processes of attention switching (in this case the N2 decline should be stronger in deviant than standard trials) or information suppression (in this case the N2 decline should be stronger in the presence, than absence, of a distractor). The obvious lack of N2b in the older group might also be related to the pronounced late fronto-central positivity of the older participants that started at the time range of P2 and that could have overlapped the N2b deflection (cf. Fig. 4). The difference between the two age groups was evident especially in the time range of P3b where younger participants showed a frontal negativity and older participants a frontal positivity. This frontality is in line with the PASA hypothesis (posterior–anterior shift with aging; [20]) holding that aging is not only associated with neural decline and decreases in brain activity, but also increasing activity reflecting functional compensation. Compensatory activations in older adults are typically found in the prefrontal cortex where activity in older adults was found to be positively correlated with performance ([20]; for review: [13]). Accordingly, in two recent speech perception studies, especially the high-performing older participants showed a pronounced frontal activation [30,29] that has been related to a mobilization of extra attentional resources. Taken together, it appeared that the two age groups used different strategies to handle the speech stimuli, with a higher inhibitory control of the young participants and a more resource-allocating strategy of the older ones. In sum, the present findings provided evidence that the processing of irregular changes of speaker position in the presence of a concurring speech stimulus resulted in a longer phase of re-orienting toward the relevant speaker in older than in younger adults. As a consequence, in everyday listening situations, snippets of a conversation can get lost and it could become more difficult for the elderly to follow the course of conversation. Disclosure statement All authors disclose no actual or potential conflicts of interest including any financial, personal, or other relationships with other people or organizations that could inappropriately influence (bias) their work. Acknowledgements The authors are grateful to Christina Hanenberg, Ines Mombrei, and Lukas Labisch for organizational assistance, and to three anonymous reviewers for valuable comments on an earlier draft of the manuscript. This work was funded by a grant from the Deutsche Forschungsgemeinschaft (DFG GE 1920/3-1). References [1] Alain C, Woods DL. Age-related changes in processing auditory stimuli during visual attention: evidence for deficits in inhibitory control and sensory memory. Psychol Aging 1999;14:507–19. [2] Amenedo E, Diaz F. Automatic and effortful processes in auditory memory reflected by event-related potentials: age related findings. Electroencephalogr Clin Neurophysiol 1998;108:361–9. [3] Anderer P, Semlitsch HV, Saletu B. Multichannel auditory event-related brain potentials: effects of normal aging on the scalp distribution of N1, P2, N2

444

[4]

[5]

[6]

[7] [8]

[9]

[10]

[11]

[12]

[13]

[14] [15] [16] [17]

[18]

[19] [20] [21]

[22]

[23]

[24] [25]

[26]

[27]

[28]

[29]

[30]

[31]

[32]

[33]

S. Getzmann et al. / Behavioural Brain Research 278 (2015) 435–445 and P300 latencies and amplitudes. Electroencephalogr Clin Neurophysiol 1996;99:458–72. Arbogast TL, Mason CR, Kidd G. The effect of spatial separation on informational and energetic masking of speech. J Acoust Soc Am 2002;112: 2086–98. Barrett G, Neshige R, Shibasaki H. Human auditory and somatosensory eventrelated potentials: effects of response condition and age. Electroencephalogr Clin Neurophysiol 1987;66:409–19. Berti S. Cognitive control after distraction: event-related brain potentials (ERPs) dissociate between different processes of attentional allocation. Psychophysiology 2008;45:608–20. Berti S. Object switching within working memory is reflected in the human event-related brain potential. Neurosci Lett 2008;28:200–5. Berti S, Grunwald M, Schröger E. Age dependent changes of distractibility and reorienting of attention revised: an event-related potential study. Brain Res 2013;1491:156–66. Bertoli S, Smurzynski J, Probst R. Temporal resolution in young and elderly subjects as measured by mismatch negativity and a psychoacoustic gap detection task. Clin Neurophysiol 2002;113:396–406. Bertoli S, Smurzynski J, Probst R. Effects of age, age-related hearing loss, and contralateral cafeteria noise on the discrimination of small frequency changes: psychoacoustic and electrophysiological measures. J Assoc Res Otolaryngol 2005;6:207–22. Brungart DS, Simpson BD. The effects of spatial separation in distance on the informational and energetic masking of a nearby speech signal. J Acoust Soc Am 2002;112:664–76. Burke DM, Shafto MA. Language and aging. In: Craik FIM, Salthouse TA, editors. The handbook of aging and cognition. New York, NY: Psychology Press; 2008. p. 373–443. Cabeza R, Dennis NA. Frontal lobes and aging: deterioration and compensation. In: Stuss DT, Knight RT, editors. Principles of frontal lobe function. New York, NY: Oxford University Press; 2013. p. 628–52. Cherry EC. Some experiments on the recognition of speech, with one and with two ears. J Acoust Soc Am 1953;25:975–9. Cooper RJ, Todd J, McGill K, Michie PT. Auditory sensory memory and the aging brain: a mismatch negativity study. Neurobiol Aging 2006;27:752–62. Czigler I, Csibra G, Ambro A. Age and information processing: event-related potential studies. Eur Psychol 1997;2:247–57. Czigler I, Csibra G, Csontos A. Age and inter-stimulus interval effects on eventrelated potentials to frequent and infrequent auditory stimuli. Biol Psychol 1992;33:195–206. Czigler I, Pató L, Poszet E, László B. Age and novelty: event-related potentials to visual stimuli within an auditory oddball—visual detection task. Int J Psychophysiol 2006;62:290–9. Darwin CJ. Listening to speech in the presence of other sounds. Philos Trans R Soc 2008;363:1011–21. Davis SW, Dennis NA, Fleck MS, Daselaar SM, Cabeza R. Que PASA?: the posterior–anterior shift in aging. Cereb Cortex 2008;18:1201–9. Escera C, Alho K, Schröger E, Winkler I. Involuntary attention and distractibility as evaluated with event-related brain potentials. Audiol Neuro-Otol 2000;5:151–66. Escera C, Yago E, Ahlo K. Electrical responses reveal the temporal dynamics of brain events during involuntary attention switching. Eur J Neurosci 2001;14:877–83. Falkenstein M, Hoormann J, Hohnsbein J. Inhibition-related ERP components: variation with modality, age, and time-on-task. J Psychophysiol 2002;16:167–75. Falkenstein M, Yordanova J, Kolev V. Effects of aging on slowing of motorresponse generation. Int J Psychophysiol 2006;59:22–9. Fitzgibbons PJ, Gordon-Salant S. Behavioural studies with aging humans: hearing sensitivity and psychophysics. In: Gordon-Salant S, Frisna RD, Popper AN, Fay RR, editors. The aging auditory system. New York, NY: Springer; 2010. p. 111–34. Folstein JR, Van Petten C. Influence of cognitive control and mismatch on the N2 component of the ERP: a review. Psychophysiology 2008;45: 152–70. Friedman D, Cycowicz YM, Gaeta H. The novelty P3: an event-related brain potential (ERP) sign of the brain’s evaluation of novelty. Neurosci Biobehav Rev 2001;25:355–73. Gaeta H, Friedman D, Ritter W, Cheng J. An event-related potential evaluation of involuntary attentional shifts in young and older adults. Psychol Aging 2001;16:55–68. Getzmann S. Handicapped due to age? Behavioral and electrophysiological correlates of speech perception of dichotically presented narratives in young and middle-aged listeners. J Psychophysiol 2012;26:132–44. Getzmann S, Falkenstein M. Understanding of spoken language under challenging listening conditions in younger and older listeners: a combined behavioural and electrophysiological study. Brain Res 2011;1415:8–22. Getzmann S, Gajewski PD, Falkenstein M. Does age increase auditory distraction? Electrophysiological correlates of high and low performance in seniors. Neurobiol Aging 2013;34:1952–62. Getzmann S, Falkenstein M, Gajewski PD. Neuro-behavioral correlates of post-deviance distraction in middle-aged and old adults. J Psychophysiol 2014;28:178–86. Gratton G, Coles MGH, Donchin E. A new method for off-line removal of ocular artifact. Electroencephalogr Clin Neurophysiol 1983;55:468–84.

[34] Grunwald T, Boutros NN, Pezer N, von OJ, Fernandez G, Schaller C, Elger CE. Neuronal substrates of sensory gating within the human brain. Biol Psychiatry 2003;53:511–9. [35] Hasher L, Zacks RT. Working memory, comprehension, and aging: a review and a new view. In: Bower GH, editor. The psychology of learning and motivation. New York, NY: Academic Press; 1988. p. 193–225. [36] Heinze HJ, Luck SJ, Mangun GR, Hillyard SA. Visual event-related potentials index focused attention within bilateral stimulus arrays I. Evidence for early selection. Electroencephalogr Clin Neurophysiol 1990;75:511–27. [37] Hillyard SA, Vogel EK, Luck SJ. Sensory gain control (amplification) as a mechanism of selective attention: electrophysiological and neuroimaging evidence. Philos Trans R Soc London Ser B 1998;353:1257–70. [38] Hoelig C, Berti S. To switch or not to switch: brain potential indices of attentional control after task-relevant and task-irrelevant changes of stimulus features. Brain Res 2010;1345:164–74. [39] Horváth J, Czigler I, Birkás E, Winkler I, Gervai J. Age-related differences in distraction and reorientation in an auditory task. Neurobiol Aging 2009;30:1157–72. [40] Horváth J, Winkler I, Bendixen A. Do N1/MMN, P3a, and RON form a strongly coupled chain reflecting the three stages of auditory distraction? Biol Psychol 2008;79:139–47. [41] Humes LE, Dubno JR, Gordon-Salant S, Lister JJ, Cacace AT, Cruickshanks KJ, Gates GA, Wilson RH, Wingfield A. Central presbycusis: a review and evaluation of the evidence. J Am Acad Audiol 2012;23:635–66. [42] Karayanidis F, Andrews S, Ward PB, Michie PT. ERP indices of auditory selective attention in aging and Parkinson’s disease. Psychophysiology 1995;32:335–50. [43] Knight RT, Scabini D. Anatomic bases of event-related potentials and their relationship to novelty detection in humans. J Clin Neurophysiol 1998;15:3–13. [44] Kok A. Age-related changes in involuntary and voluntary attention as reflected in components of the event-related potential (ERP). Biol Psychol 2000;54:107–43. [45] Kramer AF, Madden DJ. Attention. In: Craik FIM, Salthouse TA, editors. The handbook of aging and cognition. New York, NY: Psychology Press; 2008. p. 189–249. [46] Mager R, Falkenstein M, Störmer R, Brand S, Müller-Spahn F, Bullinger AH. Auditory distraction in young and middle-aged adults: a behavioral and event related potential study. J Neural Transm 2005;112:1165–76. [47] Melara RD, Rao A, Tong Y. The duality of selection: excitatory and inhibitory processes in auditory selective attention. J Exp Psychol: Hum Percept Perform 2002;28:279–306. [48] Näätänen R. The role of attention in auditory information-processing as revealed by event-related potentials and other brain measures of cognitive function. Behav Brain Sci 1990;13:201–88. [49] Näätänen R, Gaillard AWK, Mantysalo S. Early selective attention effect on evoked-potential reinterpreted. Acta Psychol 1978;42:313–29. [50] Näätänen R, Paavilainen P, Rinne T, Alho K. The mismatch negativity (MMN) in basic research of central auditory processing: a review. Clin Neurophysiol 2007;118:2544–90. [51] Näätänen R, Picton TW. The N1 wave of the human electric and magnetic response to sound: a review and an analysis of the component structure. Psychophysiology 1987;24:375–425. [52] Novak G, Ritter W, Vaughan Jr HG. Mismatch detection and the latency of temporal judgements. Psychophysiology 1992;29:398–411. [53] Patel SH, Azzam PN. Characterization of N200 and P300: selected studies of the event related potential. Int J Med Sci 2005;2:147–54. [54] Pekkonen E, Jousmaki V, Partanen J, Karhu J. Mismatch negativity area and age-related auditory memory. Electroencephalogr Clin Neurophysiol 1993;87:321–5. [55] Pekkonen E, Rinne T, Reinikainen K, Kujala T, Alho K, Naatanen R. Aging effects on auditory processing: an event-related potential study. Exp Aging Res 1996;22:171–84. [56] Polich J. Updating P300: an integrative theory of P3a and P3b. Clin Neurophysiol 2007;118:2128–48. [57] Polich J, Criado JR. Neuropsychology and neuropharmacology of the P3a and P3b. Int J Psychophysiol 2006;60:172–85. [58] Potts GF. An ERP index of task relevance evaluation of visual stimuli. Brain Cogn 2004;56:5–13. [59] Rinne T, Särkkä A, Degerman A, Schröger E, Alho K. Two separate mechanisms underlie auditory change detection and involuntary control of attention. Brain Res 2006;1077:135–43. [60] Roeber U, Widmann A, Schröger E. Auditory distraction by duration and location deviants: a behavioral and event-related potential study. Cogn Brain Res 2003;17:347–57. [61] Ruzzoli M, Pirulli C, Brignani D, Maioli C, Miniussi C. Sensory memory during physiological aging indexed by mismatch negativity (MMN). Neurobiol Aging 2012;33:625.e21–30. [62] Salthouse TA. The processing-speed theory of adult age differences in cognition. Psychol Rev 1996;103:403–28. [63] Salthouse T. A theory of cognitive aging. Amsterdam, The Netherlands: Elsevier; 2000. [64] Schröger E. A neural mechanism for involuntary attention shifts to changes in auditory stimulation. J Cogn Neurosci 1996;8:527–39. [65] Schröger E, Giard MH, Wolff C. Auditory distraction: event-related potential and behavioural indices. Clin Neurophysiol 2000;111:1450–60. [66] Schröger E, Wolff C. Attentional orienting and reorienting is indicated by human event-related brain potentials. NeuroReport 1998;9:3355–8.

S. Getzmann et al. / Behavioural Brain Research 278 (2015) 435–445 [67] Smith DB, Michalewski HJ, Brent GA, Thompson LW. Auditory averaged evoked potentials and aging: factors of stimulus, task and topography. Biol Psychol 1980;11:135–51. [68] Sussman E, Winkler I, Schröger E. Top-down control over involuntary attention switching in the auditory modality. Psychon Bull Rev 2003;10:630–7. [69] Townsend JT, Ashby FG. Stochastic modeling of elementary psychological processes. Cambridge: Cambridge University Press; 1983.

445

[70] Wascher E, Falkenstein M, Wild-Wall N. Age related strategic differences in processing irrelevant information. Neurosci Lett 2011;487: 66–9. [71] Wild-Wall N, Falkenstein M, Gajewski PD. Neural correlates of changes in a visual search task due to cognitive training. Neural Plast 2012;2012, http://dx.doi.org/10.1155/2012/529057. Article ID 529057, 11 pages.