Psychophysiology, 50 (2013), 70–73. Wiley Periodicals, Inc. Printed in the USA. Copyright © 2012 Society for Psychophysiological Research DOI: 10.1111/j.1469-8986.2012.01479.x BRIEF REPORT Separating stimulus-driven and response-related LRP components with Residue Iteration Decomposition (RIDE) BIRGIT STÜRMER,a GUANG OUYANG,b,c CHANGSONG ZHOU,b,c ANNIKA BOLDT,a,d and WERNER SOMMERa a Department of Psychology, Humboldt-Universität zu Berlin, Berlin, Germany Department of Physics, Hong Kong Baptist University, Kowloon Tong, Hong Kong c Centre for Nonlinear Studies and The Beijing–Hong Kong–Singapore Joint Centre for Nonlinear and Complex Systems, Hong Kong Baptist University, Kowloon Tong, Hong Kong d Department of Experimental Psychology, University of Oxford, Oxford, United Kingdom b Abstract When the lateralized readiness potential (LRP) is recorded in stimulus–response compatibility (SRC) tasks, two processes may overlap in the LRP, stimulus-driven response priming and activation based on response selection rules. These overlapping processes are hard to disentangle with standard analytical tools. Here, we show that Residue Iteration Decomposition (RIDE), based on latency variability, separates the overlapping LRP components from a Simon task into stimulus-driven and response-related components. SRC affected LRP amplitudes only in the stimulus-driven component, whereas LRP onsets were affected only in the response-locked component. Importantly, the compatibility effect in reaction times was more similar to the effect in the onsets of the RIDE-derived response-locked LRP component than in the unseparated LRP. Thus, RIDE-separated LRP components are devoid of distortions inherent to standard LRPs. Descriptors: ERP, Lateralized Readiness Potential (LRP), Residue Iteration Decomposition (RIDE), Stimulus– Response Compatibility (SRC) LRP in incompatible trials fits well with dual-route accounts of SRC (Kornblum, Hasbroucq, & Osman, 1990) and is seen as indicating visuo-motor priming by stimulus location via a direct route (Stürmer, Siggelkow, Dengler, & Leuthold, 2000), whereas the later correct LRP deflection is considered to reflect instructionrelated response selection (indirect route). In compatible trials, both direct visuo-motor priming and indirect instruction-related response selection activate the correct response and, therefore, result in negative LRP deflections, which overlap and seamlessly merge in the standard (composite) LRP. For compatible trials, it is therefore impossible to disentangle effects of response priming and correct response selection in the LRP. Moreover, as illustrated in Figure 1, the overlap of priming with instruction-related correct response activation distorts the onset of the correct LRPs. In compatible trials, the overlaps of both negative-going LRP components shortens the onset of the composite LRP. In incompatible trials, however, the initial positive-going activation partially cancels the overlapping negative-going activation and postpones the onset of the composite LRP. Together, both distortions exaggerate the compatibility effect in the onsets of the negative response selection-based LRPs. As a case in question, Böckler, Alpay, and Stürmer (2011) conducted a Simon task with choice responses (left vs. right hand) according to stimulus shape. Stimuli were presented above or below fixation, although stimulus location was irrelevant for response selection; response buttons were arranged within the midsaggital line. In compatible trials, stimulus location and response The lateralized readiness potential (LRP) is a useful tool indicating hand-specific response activation. The LRP is calculated from the electroencephalogram (EEG) recorded above the hand areas of the motor cortices and reflects effector-specific asymmetric brain activity. When the LRP is derived for hands, it is of negative polarity if the correct response hand is activated. In stimuluslocked LRPs (S-LRP), the onset of a negative deflection indicates the time demands of mental processes until the correct response hand is selected (Masaki, Wild-Wall, Sangals, & Sommer, 2004; Osman, Moore, & Ulrich, 1995). In stimulus-response compatibility (SRC) research, the S-LRP shows two effects. First, incompatible trials prolong the onset of the negative-going S-LRP because response selection is more time-demanding than in compatible trials. Second, in incompatible trials the correct negative-going LRP deflection is often preceded by a short-lived positive deflection indicating activation of the incorrect hand (De Jong, Liang, & Lauber, 1994; Gratton, Coles, & Donchin, 1992; Stürmer, Leuthold, Soetens, Schröter, & Sommer, 2002). The early incorrect bs_bs_banner This research was supported by the High Performance Cluster Computing Centre, Hong Kong Baptist University, and by grants of the Hong Kong Baptist University, the Hong Kong Research Grants Council (HKBU 202710) to G.O. and C.Z., the German Academic Exchange Service (DAAD) to G.O., and the German Research Foundation (STU248/3-1) to B.S. Address correspondence to: Prof. Dr. Birgit Stürmer, HumboldtUniversität zu Berlin, Institute for Psychology, Rudower Chaussee 18, 12489 Berlin. E-mail: [email protected] 70 LRP component separation by RIDE 71 Figure 1. Top panel: A schematic demonstration that the overlap of the early and late components of the LRP may advance the onset of the standard LRP for compatible trials and delay the onset for incompatible trials. LRP components related to visuo-motor priming (gray) and response selection (black) sum up to the LRP (dashed). In compatible trials (left) visuomotor priming shortens the onset of the correct negative-going LRP, whereas in incompatible trials (right), the onset is delayed. Bottom panel: (A) Standard LRP for compatible and incompatible trials (thick and thin lines, respectively) for the different arousal conditions. (B–D): LRPs computed from RIDE-derived component clusters S, C, and R, respectively, separated according to the same conditions as the standard LRP. location corresponded, whereas in incompatible trials, they did not. As expected, the onset of the correct LRP deflection was delayed in incompatible trials. In half of the trials, accessory tones preceded the Simon stimuli and shortened overall reactions times (RTs) as well as LRP onsets. Moreover, the compatibility effect was enhanced by accessory stimuli in both measures. However, the compatibility effect in LRP onsets exceeded the effect in RTs by 40 ms. It seems plausible that this excess compatibility effect is an artifact, produced by the advancement of correct LRP by the early correct activation and its delay by early incorrect activation in compatible and incompatible trials, respectively (Figure 1). Because the overlap of LRP activations via separate routes is a general problem in compatibility experiments, a separation method is highly desirable. Here, we applied Residue Iteration Decomposition (RIDE)—a method developed by Ouyang, Herzmann, Zhou, and Sommer (2011)—to the LRP. RIDE is able to separate eventrelated potential (ERP) component clusters by virtue of their latency variability. RIDE utilizes the latency variability and time markers to separate ERP components into a stimulus-locked component cluster (named S), a response-locked component cluster (named R), and an intermediate component cluster (named C). The term component cluster refers to all components with similar timelocking properties across single trials, but each may involve several ERP components. RIDE is based on the assumption that component clusters are linearly superimposed whereas their latencies vary independently, with the internal components of the same component cluster being locked to one another. By applying RIDE to LRPs in an SRC task, we aimed to disentangle the two response activations described above: (a) response activations due to visuo-motor priming via direct route processing, which should be coupled to stimulus processing and, hence, be observed in the S component separated by RIDE and (b) instruction-related response selection via the indirect route, which is closely linked to response execution and should, therefore, be observed in the R component. The present work is based on the EEG recordings reported by Böckler et al. (2011). Methods Sixteen students (mean age 22.7 years; 12 women; 15 righthanded) participated in the experiment. 72 B. Stürmer et al. Details of the stimuli and procedure can be found in Böckler et al. (2011). Compatible and incompatible Simon task conditions were orthogonally combined with three arousal conditions. A tone of 65 dB was presented either 500 or 200 ms before the visual stimulus or there could be no tone. The EEG was recorded from 64 Ag/AgCl electrodes, referenced to the left mastoid. Bandpass was 0.01 Hz to 70 Hz; sampling rate was 250 Hz. The LRP was taken from electrode sites C3 and C4 and calculated according to Coles (1989). ERP Component Decomposition with RIDE RIDE assumes that in each single EEG trial three clusters of components S, C, and R with different latencies are linearly superimposed: EEG (t ) = S (t − LS ) + C (t − LC ) + R (t − LR ) + ξ, (1) where LS, LC, and LR denote the latencies of the component clusters S, C, and R, respectively. S is assumed to be stimulus locked, so LS = 0. R is assumed to be response locked, so LR = RT. C is the component cluster in between, and its latencies have to be estimated (Ouyang et al., 2011). The time courses of S, C, and R are therefore expected to mostly represent the activations by early stimulus-related (S), intermediate (C), and response related (R) processes. RIDE obtains the waveforms of component clusters S, C, and R separately for each electrode by combining an estimate of the components from the residue of single trial EEG (EEG-ERP) and the estimate of the unknown latency Lc in an iterative manner (Ouyang et al., 2011). Each separated component had the same dimension as the ERP (1000 ms ¥ 64 channels). The core procedure of RIDE follows the work of Ouyang et al.; for a short rationale please see the introduction. Some specific modifications in detail were made for improving the performance of RIDE in the present case: 1. Data preparation. RIDE was independently and separately applied to all EEG epochs of each participant and condition, including all 64 recording sites. All single trial epochs were of 1 s duration and centered at mean amplitude. 2. Initial estimation of C latency. An initial estimation of the latency of component cluster C in single trials was made by Woody’s (1967) method separately for each recording site between 200 and 1000 ms, using the standard ERP as template. The mean latency of all channels was then taken as the overall latency of C across the scalp, denoted as LC; correspondingly, denote latency of S (stimulus onsets) as LS and latency of R (RT) as LR. 3. Residue Iteration Decomposition. With the time markers LS, LC, and LR, we used RIDE to obtain the component clusters S, C, and R, which allowed us to improve the estimate of latency LC. This procedure was iterated until LC converged and the component clusters S, C, and R and the latency LC were obtained. The baselines of stimulus-locked convolution of C and R in the interval (0–200 ms) were adjusted to zero. This operation is based on the assumption that the activity of ERP during 0–200 ms is basically stable in latency across single trials, so that the latency-variable components C and R should not contribute to 0–200 ms activity. Accordingly, the baseline of component cluster S in (0–200 ms) is readjusted to the level of standard ERP. 4. Deriving the Lateralized Readiness Potential. The LRP activations were then derived for each component cluster with the same procedure as for ERPs (Coles, 1989), subtracting the ERP derived from the ipsi- and contralateral central electrodes (C3, C4) relative to the required response hand, and averaging the difference waves. LRP Analyses After deriving the LRP for each component cluster, the singlesubject regression method with one degree of freedom by Mordkoff and Gianaros (2000) was applied to detect the onset of the standard LRP and for the LRP derived from the stimulus-synchronized component cluster R extracted by RIDE (LRP CC-R). Seven of the 164 onsets were adjusted by hand: We always assumed the S-LRP onset to occur before response execution. Two participants were completely excluded from further analyses because of their very small LRPs. Repeated measures analyses of variance (ANOVAs) were performed on RTs and LRP parameters including the factors accessory tone and compatibility. Huynh–Feldt corrections were applied if necessary, and p values were adjusted by the given e value. Results The Simon effect—calculated as the difference in mean RTs between incompatible and compatible events—was enlarged by accessory tones (no tone: 31 ms, tone at stimulus onset asynchrony [SOA] 200: 53 ms, SOA 500: 46 ms) in the Böckler et al. (2011) study. Likewise, the Simon effect in standard LRP onsets was numerically increased by tones (no tone: 67 ms, tone at SOA 200: 106 ms, tone at SOA 500: 111 ms). However, only the main effect of compatibility was significant in S-LRP onsets, F(1,13) = 59.63, p < .001, partial h2 = .821. No interaction of compatibility and accessory tones showed up in S-LRP onsets, F(2,26) = 1.96, p = .16, partial h2 = .123. Figure 1d, bottom, shows the LRP for the RIDE-derived stimulus synchronized component cluster R (LRP CC-R). The onsets of these waveforms showed main effects of compatibility (Mc = 208 ms, Mic = 257 ms); F(1,13) = 14.09, p < .01, partial h2 = .520, but not of accessory tone (M for no tone, tone at SOA 200 and 500 ms = 243, 223, and 233 ms, respectively); F(2,26) = 1.35, p = .277, partial h2 = .094. The interaction of compatibility and accessory tone was far from significant; F < 1, partial h2 = .002. Figure 1b, bottom, shows the LRPs for the RIDE-derived component cluster S (LRP CC-S), and Figure 1c, bottom, shows the LRP for the component cluster C (LRP CC-C). Overall, the LRP CC-C does not contribute to present LRP effects. Please note that this only refers to the hand-related asymmetry that is extracted by the LRP calculation and only means that there is no such asymmetry in the component cluster C. However, the LRP CC-S beautifully captures the early visuo-motor priming-related LRP activation with positive- and negative-going deflections of similar magnitude, clearly influenced by the accessory tones. One may have noticed that these positive- and negative-going deflections are not symmetric relative to the x-axis—as implied by the model in Figure 1—but they are symmetric relative to a negative-going slope of the S-LRP. Such slopes due to a nonreturn to baseline are to be seen in most LRP studies (e.g., Gratton et al., 1992; Stürmer et al., 2002) but, to the best of our knowledge, have not found scientific attention. RIDE allocates the slope, which has no significant latency jitter within our 1-s epoch, to component cluster S. The earlier peak in the CC-S LRP from the incompatible as compared to the compatible condition is a consequence of the negative-going slope: A negative slope superimposed on a positive LRP component separation by RIDE 73 wave will shift the maximum of the composite signal toward earlier latencies, whereas superposition on a negative wave will delay the latency of the minimum. A conceivable way of dealing with the slope problem would be a detrending of the single trials prior to RIDE. However, for the moment, we preferred to retain this trend because it is a property of most LRPs in the literature. An ANOVA on the average LRP CC-S amplitude within the time interval 200–280 ms after Simon stimulus onset resulted in a main effect of compatibility, F(1,13) = 41.49, p < .001. Single comparisons showed that the compatible no-tone condition significantly differed from both compatible tone conditions, Fs(1,13) > 5.03, ps < .043. Incompatible conditions differed significantly between the no-tone and tone 200 conditions, F(1,13) = 7.60, p < .016. Therefore, the S component replicated all amplitude effects of compatibility and arousal in the standard ERP. Discussion We assessed the feasibility of separating the early priming-based LRP component from the later response-selection based component by means of RIDE, a method for ERP component separation based on latency variability. As expected, RIDE yielded two LRP components, one extracted from component cluster S (LRP CC-S) and one extracted from the component cluster R (LRP CC-R). No significant LRP was seen on component cluster C. Compared to the traditional ERP method, RIDE provides several novel insights. The LRP CC-S captures the direct route of early visuo-motor processing and fit with standard Simon effect results in the LRP amplitude. In incompatible trials, the LRP CC-S showed a clear positive deflection after 200 ms, which later on shifted toward more negative values. In compatible trials, no such positive deflection was present in the early part of the LRP CC-S. The LRP CC-R is devoid of early activation and displays a significant Simon effect in the onset (M = 49 ms). Importantly, this LRP CC-R Simon effect was very close to that found in mean RT (M = 43 ms), whereas in the standard LRP the mean effect had been 95 ms. This result nicely supports the hypothesis that the overlap of visuo-motor priming-related LRP and response selection-related LRP advances and delays the onsets of correct LRP activations in compatible and incompatible trials, respectively. In addition, the negative deflection in the LRP CC-S of compatible trials and the positivity in the incompatible conditions (see Figure 1a,b) confirm the postulated overlap of the mechanisms of visuo-motor priming and response selection. 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Medical and Biological Engineering and Computing, 5, 539–554. (Received May 21, 2012; Accepted September 5, 2012)
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