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Original Research ARTICLE
Both attention and prediction are necessary for adaptive neuronal tuning in sensory processing
- 1Université Paris Descartes, Sorbonne Paris Cité, Paris, France
- 2CNRS, Laboratoire Psychologie de la Perception, UMR 8242, Paris, France
- 3Department of Psychology, University of Jyväskylä, Jyväskylä, Finland
The brain as a proactive system processes sensory information under the top-down influence of attention and prediction. However, the relation between attention and prediction remains undetermined given the conflation of these two mechanisms in the literature. To evaluate whether attention and prediction are dependent of each other, and if so, how these two top-down mechanisms may interact in sensory processing, we orthogonally manipulated attention and prediction in a target detection task. Participants were instructed to pay attention to one of two interleaved stimulus streams of predictable/unpredictable tone frequency. We found that attention and prediction interacted on the amplitude of the N1 ERP component. The N1 amplitude in the attended/predictable condition was larger than that in any of the other conditions. Dipole source localization analysis showed that the effect came from the activation in bilateral auditory areas. No significant effect was found in the P2 time window. Our results suggest that attention and prediction are dependent of each other. While attention might determine the overall cortical responsiveness to stimuli when prediction is involved, prediction might provide an anchor for the modulation of the synaptic input strengths which needs to be operated on the basis of attention.
Recent theories of sensory processing consider the brain as a proactive system which adapts quickly to the environment. Neurons in the sensory cortices can undergo short-term, task-dependent, and context-specific changes in receptive field properties when attention and prediction are involved (Fritz et al., 2003, 2007, 2008). Such adaptive plasticity driven by attention and prediction can be the underlying mechanism for the optimization ofperception.
Attention is suggested to have a global effect on perception at an early stage of sensory processing. Electroencephalography (EEG) studies revealed the neuronal consequences of attention on event-related potentials (ERPs), particularly the enhancement of the N1 (Hillyard et al., 1973; Alcaini et al., 1994; Lange et al., 2003, 2006; Lange and Röder, 2006). This may result from changes in the selectivity of neurons in the sensory cortex (Chawla et al., 1999; Kastner et al., 1999; Ahveninen et al., 2006). Specifically, research showing that the auditory N1 response is modulated by task demands and notched-noise masking suggests that the spectrotemporal receptive fields of neurons are tuned according to attentional manipulations (Kauramäki et al., 2007), as attention excites neurons responsive to attended features and inhibits neurons responsive to unattended features (Fritz et al., 2003, 2007, 2008; Jääskeläinen et al., 2007). Neurocomputational studies demonstrated that attention may function via optimizing the synaptic gain to represent the precision of sensory information during hierarchical inference (Feldman and Friston, 2010).
Prediction, or the statistical regularity in the environment, is also suggested to modulate the early stage of sensory processing, albeit its effect on ERPs manifests as a suppression of the N1 (Lange, 2013). Prediction-related N1 suppression was demonstrated when participants had foreknowledge of the upcoming stimuli (Schafer and Marcus, 1973; Schafer et al., 1981; Lange, 2009; SanMiguel et al., 2013; Timm et al., 2013). The predictive coding model postulates that the prediction effect indexes the difference in neurocomputational demand for predictable/unpredictable information (Friston, 2005; Egner et al., 2010). Specifically, it indexes how much of the sensory input cannot be accounted for by the internal model. Moreover, prediction was reported to alter the contrast gain of sensory evidence accumulation (Melloni et al., 2011; Rohenkohl et al., 2012). Neurophysiologically, this is reflected in sharper sensory representations where the reduction of neuronal activity is smaller in neurons responsive to predictable features than in neurons responsive to unpredictable features (Kok et al., 2012a).
Despite their ERP effects being opposite, the relation of attention and prediction remains undetermined. This might be due to the conflation of these two mechanisms in the literature, where attention and prediction were often treated as the same concept (Corbetta and Shulman, 2002). However, attention and prediction can rely on orthogonal sources of information (Summerfield and Egner, 2013). While attention operates on the basis of motivational relevance, prediction operates on the basis of prior likelihood (Summerfield and Egner, 2009). It is possible that attention and prediction are two independent mechanisms which may have antagonistic (Summerfield and Egner, 2009) or additive (Timm et al., 2013) effects on neuronal signals for sensory processing. Alternatively, attention and prediction may be dependent of each other. One possibility is that one of the two mechanisms is necessary for the other to take effect, but not the other way round (Kok et al., 2012b). Another possibility is that such dependency is bidirectional, with both attention and prediction being necessary to modulate sensory processing.
To examine the relation between these two top-down mechanisms, we orthogonally manipulated attention and prediction in a target detection task. Participants were instructed to pay attention to one of two interleaved stimulus streams of predictable/unpredictable tone frequency. Using EEG, we quantified N1 and P2 as dependent variables given that the former is involved in auditory perception and the latter is suggested to reflect the comparison between the sensory input and the internal model (Evans and Federmeier, 2007; Costa-Faidella et al., 2011). The design allowed us to evaluate whether attention and prediction are dependent of each other, and if so, how these two top-down mechanisms may interact on sensory processing.
Materials and Methods
Sixteen healthy volunteers (average age 28; six males; all right-handed) with no history of neurological, psychiatric, or hearing impairments as indicated by self-report participated in the experiment. Participants gave written informed consent and were paid for participation. Ethical approval was granted by the Comité de Protection des Personnes (CPP) Ile de France II. The experiment conforms with The Code of Ethics of the World Medical Association (Declaration of Helsinki).
Sinusoidal tones with a loudness of 80 phons (i.e., 80 dB for tones of 1000 Hz) were generated using Matlab. The duration of each tone was 50 ms (including 5 ms rise/fall times). The frequency of each tone was within the range of 261.626–493.883 Hz and 2093.000–3951.070 Hz, matching the absolute frequency of two sets of seven natural keys on a modern piano (low frequency set: C4 D4 E4 F4 G4 A4 B4; high frequency set: C7 D7 E7 F7 G7 A7 B7). Within each frequency set, the predictable/unpredictable stimulus streams were created. The predictable stimulus stream consisted of 400 pairs of tones. Each pair of tones was arranged in ascending order with the second tone being two natural keys higher than the first tone (e.g., C4-E4; G7-B7). The unpredictable stimulus stream consisted of 400 pairs of tones. Each pair of tones was arranged in random order with the second tone being any tone except the repetition of the first tone (e.g., C4-F4; G7-F7). The tones in each condition were of equal variability in frequency.
The predictable/unpredictable stimulus streams from different frequency sets were interleaved to allow for the efficient manipulation of attention on the two stimulus sets. To counterbalance the mapping between predictable/unpredictable stimulus streams and high/low frequency sets, half of the participants were presented with a low-frequency predictable stimulus stream interleaved with a high-frequency unpredictable stimulus stream and half of the participants were presented with a high-frequency predictable stimulus stream interleaved with a low-frequency unpredictable stimulus stream. To counterbalance the sequential relation of the predictable/unpredictable stimulus streams, half of the blocks started with the predictable stimulus stream and half of the blocks started with the unpredictable stimulus stream. A stimulus onset asynchrony (SOA) of 500 ms was used (Figure 1). 10% of the tones in one of the two stimulus streams served as targets attenuated in loudness by 20 dB, independent of whether they were the first/second tones in each pair of tones. Stimulation was created individually for each participant.
FIGURE 1. Experimental design. The predictable/unpredictable stimulus streams from different frequency sets were interleaved. The numbers indicate the tones being the first/second tones in each pairs of tones. Unaware of the manipulation of prediction on tone frequency, participants were instructed to pay attention to one of the stimulus streams in different blocks where tones of attenuated loudness may appear.
Participants were presented with a total of eight blocks of 100 pairs of tones via headphones (Sennheiser PX200), with each block including 50 predictable pairs of tones and 50 unpredictable pairs of tones. Unaware of the manipulation of prediction, participants were instructed to pay attention to one of the stimulus streams (i.e., high/low frequency) in different blocks where tones of attenuated loudness may appear. To monitor whether participants followed the instructions correctly, participants were required to press a key when they detected a softer tone, which randomly occurred 10 times in each block. No practice session was provided. Block order was counterbalanced across participants. The experiment was administered conjointly with another EEG experiment which is to be reported elsewhere.
The stimuli of interest were the second tones in each pair of tones, which can be attended/predictable, attended/unpredictable, unattended/predictable, and unattended/unpredictable. Note that the manipulations of attention and prediction were both introduced on the basis of tone frequency. Moreover, all stimuli were presented binaurally. Therefore, there was no spatial effect in the current study.
Data Recording and Analysis
EEG recording and pre-processing
EEG was recorded with 64 active electrodes (actiCAP, Brain Products GmbH, Germany) conforming to the international 10–10 system. The sampling rate was 500 Hz. No online/offline filter was used. The data was recomputed to average reference. Target stimuli and the first stimuli following target stimuli were removed. Epochs extended from -100 to 500 ms relative to stimulus onset, using a 100 ms pre-stimulus baseline. Ocular artifact correction was conducted with independent component analysis in EEGlab (Delorme and Makeig, 2004). Epochs containing voltage deviations exceeding ±80 μV relative to baseline at any of the electrodes were rejected. The trial numbers after artifact rejection in each condition are listed in Table 1.
ERP analysis
ERP analysis was based on a temporal principal component analysis (PCA) in SPSS 20. The temporal PCA statistically decomposes the ERP waveforms into constituent building blocks, which affords objective data-driven ERP component measures when compared to the conventional peak-picking methods (Kayser and Tenke, 2003; Dien et al., 2005; Kayser and Tenke, 2006; Dien, 2012). Moreover, it is not susceptible to the influences of high-frequency noises and low-frequency drifts in the data as the conventional peak-picking methods (Luck, 2005). Covariance matrix and Promax rotation were used. All components accounting for a total of 99% of the variance (maximum iterations for convergence = 500) were included in the rotation (Promax Kappa = 4). The temporal decomposition provided a set of component loadings reflecting the contribution of each time point on the temporal principal components. The component loadings were used to derive component scores which represent the magnitude of neural activity within the time windows of interest. We identified N1 and P2 by selecting components showing typical N1 and P2 latency and topography. Electrodes showing the largest N1 component score (i.e., F2) and P2 component score (i.e., Cz) were considered as representative electrodes for the components of interest. Therefore, their component scores were used as input for the 2 (attended/unattended) × 2 (predictable/unpredictable) repeated measures analysis of variance (ANOVA).
Behavioural Data
Overall, participants' performance in the target detection task was at ceiling (Hit: mean = 0.96, SD = 0.03; False alarm: mean < 0.01, SD < 0.01; RT: mean = 539.34, SD = 62.21), confirming that participants followed the instructions correctly. The ceiling performance rendered it unlikely that participants' attention alternated between the two stimulus streams within blocks, which would have brought down participants' performance. There was no difference between participants' performance when they attended to predictable/unpredictable stimulus stream [Hit: t(15) = 0.30, p = 0.77; False alarm: t(15) < 0.01, p = 1.00; RT: t(15) = 0.32, p = 0.75]. The equivalent performance across blocks excluded the possibility that task difficulty may be a confounding factor in the current study.
ERP Data
Figure 2 shows the grand average ERPs on nine representative electrodes (F3, Fz, F4, C3, Cz, C4, P3, Pz, P4). N1 with a frontocentral distribution and P2 with a central distribution are evident. Figure 3 shows the component loadings of 111 components in the temporal PCA. The temporal PCA yielded components clearly corresponding to N1 and P2 in the grand average ERPs. Figure 4 shows the N1 and P2 component scores on electrodes showing the largest responses (N1: F2; P2: Cz) and the topographical distribution of the components in each condition.
FIGURE 2. Grand average ERPs on nine representative electrodes (F3, Fz, F4, C3, Cz, C4, P3, Pz, P4) lowpass filtered at 30 Hz for visual presentation purposes. N1 and P2 in the grand average ERPs are denoted.
FIGURE 3. Component loadings of 111 components in the temporal PCA. Components clearly corresponding to N1 and P2 in the grand average ERPs are marked with thick lines.
FIGURE 4. The N1 and P2 component scores on electrodes showing the largest responses (N1: F2; P2: Cz) and the topographical distribution of components in each condition. Error bars depict one standard error above and below the mean.
The 2 × 2 repeated measures ANOVA showed a significant main effect of attention with attended stimuli triggering enhanced activity compared to unattended stimuli [F(1,15) = 6.50, p < 0.05] and a marginally significant main effect of prediction with predictable stimuli triggering enhanced activity compared to unpredictable stimuli [F(1,15) = 3.40, p < 0.10]. More importantly, there was a significant attention × prediction interaction [F(1,15) = 4.61, p < 0.05]. Post hoc comparisons showed that the component score was larger for attended stimuli than for unattended stimuli in the predictable condition [t(15) = -3.42, p < 0.01] but not in the unpredictable condition [t(15) = -0.60, p = 0.56] and that the component score was larger for predictable stimuli than for unpredictable stimuli in the attended condition [t(15) = -2.40, p < 0.05] but not in the unattended condition [t(15) = -0.63, p = 0.54]. Moreover, the component score for attended/predictable stimuli was larger than that for unattended/unpredictable stimuli [t(15) = -2.44, p < 0.05]. In other words, the component score in the attended/predictable condition was larger than that in any of the other conditions. To examine the neural origin of the effects of attention and prediction, the component scores on all electrodes for N1 were further imported into BESA Research 6.0 for dipole source localization (Scherg and von Cramon, 1986). A four-shell ellipsoidal head model was used. Two equivalent current dipoles were randomly seeded and freely fitted. The analysis showed that all four conditions had sources close to each other in the bilateral auditory areas (Figure 5). The average Talairach coordinates across the four conditions (left, x: -40.5, y: -20.9, z: 6.5; right, x: 37.0, y: -18.9, z: 13.5) are close to the approximate location of the Heschl's gyrus (e.g., the average limits in Penhune et al., 1996 are for the left x: -32.5 to -62.3, y: -33.2 to -4.9, z: 1.4 to 14.2 and for the right x: 34.7 to 58.2, y: -32.8 to -4.7, z: 1.1 to 17.7). The two dipole model had residual variances of 1.22% (attended/predictable), 1.62% (attended/unpredictable), 2.51% (unattended/predictable), and 2.45% (unattended/unpredictable). The result suggests that the effects of attention and prediction are based on the activation of neuronal populations in the bilateral auditory areas.
FIGURE 5. Source locations of the N1 response in each condition based on the topographical distribution of the component in the temporal PCA.
In the 2 × 2 repeated measures ANOVA, neither the main effect of attention [F(1,15) = 0.44, p = 0.52] nor the main effect of prediction [F(1,15) = 0.70, p = 0.42] was significant. The attention × prediction interaction was marginally significant [F(1,15) = 4.15, p = 0.06]. However, there was no significant effect in Tukey LSD post hoc comparisons (Attention effect in predictable condition: t(15) = -1.99, p = 0.07; Attention effect in unpredictable condition: t(15) = 1.08, p = 0.30; Prediction effect in attended condition: t(15) = -0.61, p = 0.55; Prediction effect in unattended condition: t(15) = 1.84, p = 0.09).
The orthogonal design in the current study allowed us to evaluate whether attention and prediction are dependent of each other, and if so, how these two top-down mechanisms may interact for the optimization of perception. We found that attention and prediction interacted on the amplitude of the N1 ERP component generated in the auditory cortices. The N1 amplitude in the attended/predictable condition was larger than that in any of the other conditions. The relatively early latency of the interaction between these two variables is in line with the idea that attention and prediction can synergistically enhance perceptual analysis through top-down pathways (Doherty et al., 2005; Chaumon et al., 2008). Furthermore, it seems that attention and prediction rely on each other to form a tonically maintained set which can selectively include/exclude sensory input for further processing (Hillyard et al., 1973; Kok et al., 2012b). In other words, both attention and prediction are involved in the optimization of sensory processing.
Precious research reported that neurons in the sensory cortices can undergo short-term, task-dependent, and context-specific changes in the receptive field properties (Fritz et al., 2003, 2007, 2008). Such changes in the selectivity of sensory cortex neurons are believed to be the mechanism underlying the optimization of perception (Ahveninen et al., 2006; Kauramäki et al., 2007; Kok et al., 2012a). Extending the idea, our results further suggest that both attention and prediction are needed for such rapid plasticity to occur. Specifically, one may speculate that while attention determines the overall cortical responsiveness to stimuli (Lakatos et al., 2008; Busch et al., 2009; Schroeder and Lakatos, 2009), prediction serves as an anchor for the selective modulation of synaptic input strengths. An interesting question for future research is whether it is possible to disentangle the synergistic neuronal effects of attention and prediction on sensory processing.
The prediction-dependent attention effect demonstrates, for the first time, that attention alone may not be able to reshape perceptual inferences. It is unknown whether the prediction participants formed in the current study was implicit or explicit. However, it seems that prediction, be it implicit or explicit, is needed for attention effects to occur. At first glance, this idea seems difficult to reconcile with previous studies showing that attention can selectively modulate the response of neuronal subpopulations that prefer the attended stimulus features (Martinez-Trujillo and Treue, 2004; Liu et al., 2007). However, it is noteworthy that, in previous studies, attended stimulus features were also predictable stimulus features, making it difficult to disentangle the influence of prediction on attention effects. Our finding of the prediction-dependent attention effect suggests that prediction may be an important construct for attention.
The attention-dependent prediction effect, on the other hand, sheds light on the dynamic influence of attention on prediction. Notably, using functional magnetic resonance imaging (fMRI) and visual stimuli, Kok et al. (2012b) reported an approach of manipulating attention and prediction that is similar to ours. In the attended condition, our finding of prediction-related enhancement of the N1 amplitude from the auditory cortices was consistent with Kok et al.'s finding of prediction-related enhancement of activation in early visual cortex. Such enhancement of neuronal responses confirm the notion that the engagement of attention may increase the weighting of sensory information, resulting in enhanced neuronal responses to predictable stimuli relative to unpredictable stimuli (Rao, 2005). The enhanced N1 is nicely in line with the predictive coding theory of sensory processing. The predictive coding theory suggests that perception entails the propagation of signals from two distinct types of neuronal populations, the representation neurons, and the prediction error neurons (Friston, 2005, 2009; Egner et al., 2010). Although there was some debate in recent neurocomputational research concerning which type of neurons is modulated by attention (Spratling, 2008, 2010; Feldman and Friston, 2010), it was shown that both models presume enhanced responses to predictable stimuli relative to unpredictable stimuli in the attended condition. In contrast, in the unattended condition, our null result was inconsistent with Kok et al.'s finding of prediction-related suppression of activation in early visual cortex. While caution should be taken for the interpretation of our null result, we speculate that the discrepancy may be related to the different manipulations of attention and prediction.
Concerning the manipulation of attention, the discrepancy may be explained by a competition hypothesis borrowed from research on mismatch negativity (MMN). Mismatch negativity is generated to violations of abstract stimulus prediction rules (Tervaniemi et al., 1994; Paavilainen et al., 1999), reflecting the detection of deviant stimuli independent of attention (Näätänen, 1990; Näätänen et al., 2007). Importantly, a competition hypothesis suggests that, although attention does not affect the deviant detection process per se, it may influence the sensory information reaching the deviant detection process (Sussman et al., 2003). For example, when certain stimulus features are attended in a competing stimulus stream, the detection of stimuli deviating in certain stimulus features can be accordingly abolished in the unattended channel, leading to the absence of the MMN.
We suggest that whether the prediction effect appears in the unattended condition depends on the degree of such competition for cognitive resources. In the current study, attention was manipulated in the "filtering" manner. In this case, participants may inhibit the processing of unattended stimuli stream at an early stage to get rid of unnecessary information. In the study of Kok et al. (2012b), on the contrary, attention was manipulated in the "probabilistic" manner. In this case, participants may not inhibit the processing of unattended stimuli until the level concerning behavioral output. Consequently, the competition for cognitive recourses may be too weak to eliminate the possibility that unpredictable stimuli automatically attract attention, resulting in enhanced neuronal responses (Larsson and Smith, 2012). The same line of argumentation holds for previous studies demonstrating prediction-related suppression effects, in which participants' attention was either not controlled (Schafer and Marcus, 1973; Stefanics et al., 2010; Besle et al., 2011) or directed away from the manipulation of prediction using a cover task (Schafer et al., 1981; Summerfield et al., 2008; Lange, 2009; Todorovic et al., 2011; Kok et al., 2012a; Todorovic and de Lange, 2012). Given that participants' attention still stayed focused on the stimuli in these tasks, it is difficult to exclude the possibility that the observed prediction-related suppression effects reflected the enhanced neuronal responses to unpredictable stimuli.
Alternatively, whether the prediction effect appears in the unattended condition may depend on the particular manipulation of prediction. In the current study, the unpredictable condition consisted of stimuli randomly selected from a given frequency set. In other words, it is difficult for participants to form a specific prediction about the frequency of the upcoming stimuli in the first place. Therefore, the activation in sensory cortex may reflect exclusively the neuronal signals triggered by the presence of unpredictable stimuli. In the study of Kok et al. (2012b), on the contrary, the unpredictable condition consisted of stimuli which were presented as oddballs that violated participants' prediction. Consequently, the activation in sensory cortex may reflect not only the neuronal signals triggered by the presence of unpredictable stimuli but also the neuronal signals triggered by the absence of predictable stimuli. Such increase in the neuronal signals in the unpredictable condition may be the reason why prediction-related suppression can be observed.
On the other hand, no significant effect was found in the P2 time window. At best, there was a marginally significant interaction between attention and prediction. While caution should be taken in interpreting the results, the pattern of the interaction suggests that attention reversed the direction of prediction effects on the P2. In the attended condition, predictable stimuli suppressed the P2. In the unattended condition, predictable stimuli enhanced the P2. Interestingly, the pattern is opposite to the findings of Kok et al. (2012b), where prediction-related hemodynamic responses in early visual cortex were enhanced when the stimuli were attended and suppressed when the stimuli were unattended. More research is needed to examine whether the interaction between attention and prediction may change over time and how such dynamics manifest in hemodynamic responses.
Overall, our findings suggest that future research on attention and its relation to prediction needs to differentiate between different manipulations of attention and prediction. Moreover, our results confirm the importance of incorporating the modulatory effect of attention in the predictive coding theory. The predictive coding theory postulates that the prediction effect indexes the difference in neurocomputational demand for predictable/unpredictable information (Friston, 2005; Egner et al., 2010). Specifically, it is suggested that predictable information, relative to unpredictable information, elicits smaller prediction error signals. The smaller prediction error signals are associated with reduced connectivity within a hierarchical cortical network (Garrido et al., 2007, 2009), which accounts for the observations of prediction-related suppression in evoked responses (Wacongne et al., 2011). The predictive coding theory seems contradictory to our findings of significant prediction-related N1 enhancement in the attended condition and the lack of prediction-related N1 suppression in the unattended condition, unless the modulatory effect of attention is taken into consideration. Recent computational models demonstrated that attention emerges naturally in a similar Bayes-optimal scheme as the inference about the precision of the causes of sensory input (Friston, 2009; Feldman and Friston, 2010). It is, thus, possible that the prediction effect does not reflect merely the different prediction error signals. Rather, it reflects how the precision of the causes of sensory input is weighed against the prediction error signals. In other words, it is the attention effect, rather than the prediction effect, that is dominant here. In the attended condition, the precision of the causes of sensory input increases. The increased precision may manifest as an imaginary tuning curve with narrower bandwidth. Moreover, it may outweigh the prediction error signals to predictable input over the prediction error signals to unpredictable input. This is when the prediction-related enhancement can be observed. In the unattended condition, on the other hand, the precision of the causes of sensory input decreases. It can manifest as an imaginary tuning curve with wider bandwidth. In this case, the amount of the decrease in precision may determine to what extent the prediction error signals to unpredictable input can be canceled out. The larger the decrease in precision, the more it cancels out the prediction error signals to unpredictable input. At some point, the prediction error signals to unpredictable input would become really small, rendering the prediction-related suppression negligible. A real life analogy is that, when one pays minimum amount of attention to the stimuli, the boundary between the unpredictable input and the predictable input is of little importance. This mechanism could be tested in detail in future research.
Conflict of Interest Statement
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
This research was supported by a grant from the Agence Nationale de la Recherche (INTACT ANR-09-BLAN-0318). We thank Trevor Agus for help on stimulus calibration and the Paris Descartes Platform for Sensorimotor Studies (Université Paris Descartes, CNRS, INSERM, Région Ile-de-France) for supporting the experimental work presented here.
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Kayser, J., and Tenke, C. E. (2003). Optimizing PCA methodology for ERP component identification and measurement: theoretical rationale and empirical evaluation. Clin. Neurophysiol. 114, 2307–2325. doi: 10.1016/S1388-2457(03)00241-4
Kayser, J., and Tenke, C. E. (2006). Consensus on PCA for ERP data, and sensibility of unrestricted solutions. Clin. Neurophysiol. 117, 703–707. doi: 10.1016/j.clinph.2005.11.015
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Keywords: attention, prediction, sensory processing, electroencephalography, event-related potentials
Citation: Hsu Y-F, Hämäläinen JA and Waszak F (2014) Both attention and prediction are necessary for adaptive neuronal tuning in sensory processing. Front. Hum. Neurosci. 8:152. doi: 10.3389/fnhum.2014.00152
Received: 22 January 2014; Accepted: 28 February 2014;
Published online: 26 March 2014.
Edited by:Harriet Brown, University College London, UK
Reviewed by:Ana Todorovic, Radboud University Nijmegen, Netherlands
Ryszard Auksztulewicz, University College London, UK
Copyright © 2014 Hsu, Hämäläinen and Waszak. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.
*Correspondence: Yi-Fang Hsu, Université Paris Descartes, Sorbonne Paris Cité and CNRS, Laboratoire sychologie de la Perception, UMR 8242, 45 Rue des Saints-Pères, 75006 Paris, France e-mail: firstname.lastname@example.com


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Even as advisors jump ship, top execs are staying put
Top industry talent is staying put — for now.
Only 10% of high-ranking executives from wealth management, asset management and fintech companies reported moving to a new firm last year, according to an annual survey of talent trends by Kathy Freeman Company, a California-based executive search firm.
That's the lowest number reported in the past seven years, when the percentage of executives surveyed switching jobs ranged from 22% in 2010 to 15% in 2015.
Professional satisfaction appears to be greater at the top. While executives are mostly staying put, more than half of lower-ranked financial advisors have either switched firms in the last five years or are seriously considering a move, according to a recent Fidelity study.
"Life is good for top executives," said Kathy Freeman, principal of the eponymous firm. "People are getting paid well and if there's no pain point, why move? Why blow a good thing?"
Indeed, 49% of respondents said their total pay increased last year, and an equal number expect their compensation to rise again in 2018.
But that could change quickly if there's a bear market. "Compensation will inevitably come down and people will start looking around," Freeman says.
In the current environment, however, firms looking to attract top talent will have to go the extra mile.
"When considering a change, a candidate's key concern is always the sustainability of the firm," according to the report, titled "Late Adopters Finish Last: Why Change is Critical in Today's Talent Market." "Potential candidates will be looking for reassurance that they won't get caught in a last-hired, first-hired scenario if the market has a sizable correction."
Assuming compensation meets competitive standards, nearly half of the executives surveyed said a continuous challenge in their current role was the main reason they haven't moved. "It's not just about money," says Freeman. "In a market where pay is trending up, competitive compensation is a given rather than a meaningful motivator for a transition."
"Ask candidates to help frame out what the job can be," says recruiter Kathy Freeman.
Firms seeking to lure department heads and corporate leaders should consider having a broader framework for responsibilities when recruiting established executives, Freeman says.
"That level of talent is interested in opportunities where they can be creative," she explains. "Let them help fill in the blanks when discussing the parameters of a new position. Ask candidates to help frame out what the job can be. They want to see examples of fresh thinking."
All firms, including those who want to retain top talent, should review existing compensation models, the report recommends.
The industry is beginning to see structural changes in compensation, Freeman says, with more to come. Indeed, over one-third of executives surveyed anticipated a change in compensation structure this year. Legacy compensation packages may no longer be sustainable, the report warns.
Some firms are beginning to pay a higher base salary because variable compensation can be so unpredictable, Freeman says. "We're seeing more specificity around what firms pay so there are no surprises at the end of the year," she notes.

subset: commoncrawl


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Also meant to ask - I looked through the webinar videos but did not see an AP best practices demo. Is one available and if not, maybe a good one to do for the future?
Currently, our AP process is as follows:
This current process works okay and there are checks and balances in place, but we run into issues such as:
Thanks for any suggestions.


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Abandoned Baby Horse Got Rescued, Now He Can't Sleep Without His Teddy Bear 2k
Breeze the baby pony had a struck of luck when he was found and rescued by the good people from the local equine sanctuary. He was found alone and scared at the Dartmoor National Park, merely hours after his birth. If he wasn't found for another few hours, chances are he would not have been here today. Now the one-week-old foal loves to sleep on the lap of a giant teddy bear, which acts as a surrogate mom to help the foal feel comfortable and protected.


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This five bedroom house is a stunning property, ideal for students looking for a high standard of accommodation. Situated within the Eaton area, means all amenities are close by.
The property comes fully furnished, with brand new pine furniture on all of the 5 bedrooms. The kitchen is fully equipped with a fridge-freezer, washing machine and microwave. There is two bathrooms within the property, one which is fitted with a shower over the bath and the other has a separate shower and bath. Shared living room and neat and tidy low maintenance front and rear garden. Gas central heating.
This property is well equipped and furnished to a high standard. There is off-road parking and the property is conveniently located for supermarkets and food outlets with great transport links to Norwich City Centre and The University Hospital.
Property is available as a group rental or individual rooms. Both single and double rooms available.


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Over Christmas break I dusted off a book of poems by a Norwegian writer named Olav Haugue—a book I've owned for years but never read cover to cover. What a serendipitous move on my part. I came across a number of tiny but illuminating poems, including this one:
Don't Come to Me with the Entire Truth
Don't come to me with the entire truth.
Don't bring the ocean if I feel thirsty,
Nor heaven if I ask for light;
But bring a hint, some dew, a particle,
As birds carry drops away from a lake,
And the wind a grain of salt.
(translated by Robert Bly)
We might consider this a revision, or at least an updating, of Dickinson's savvy advice: "Tell all the truth but tell it slant— Success in Circuit lies." She ends her own thimble-sized poem with these plain but oracular words: "The Truth must dazzle gradually / Or every man be blind."
What does this have to do with your studies? Too often we're waiting for some grand truth, some scintillating conversation, some giant, about-face paradigm shift to rattle our bones when something small will do. As you settle into new classes, with their constellation of fresh readings and classmates and (cross your fingers) remarkable professors, cultivate the ability to be moved in the smallest of ways. Nibble, experiment, explore. And let this new alertness leaven all aspects of your life. Who knows, perhaps that thing you most need is already within your grasp, as it was with me: let me see, a little to the right, one shelf up, huh, I wonder what's inside that forgettable cover?
Lance Larsen, Chair
Department of English


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Purposeful Placement
Advertising is a world of its own, but BrightBox believes in firmly integrating your advertising efforts within your larger marketing plan. This way, advertising is properly listed as a tactic in your monthly strategy, utilized and integrated with every other touch point your customer experiences. There are many reasons to advertise in various print and digital mediums, but more often than not it's for one of two purposes--increasing brand awareness or specifically promoting a product or service of greater interest. The BrightBox team offers you unmatched advertising relationships. From years of close contact with both local and national industry-specific websites and publications, we leverage relationships for your good--bigger and better opportunities to showcase your brand and promote your products and services. There are never-ending possibilities, but with our team, you get a recommended ad spend for media buys that put your brand in the right places, at the right times.
Print & Digital
Looking at advertising opportunities, our team selects the right ad packages that provide you with the best exposure and ROI in mind. Results-driven, BrightBox manages your media buy spending and gives you peace of mind from the hassle of putting together multiple touch points, tracking ad success and managing ad submissions month to month.
Traditional print advertising is anything but obsolete. In fact, for many industries, print advertising is still demanded by consumers as the preferred form of communication on promotions and specials. For other industries, print advertising is simply keeping up with the status quo, ensuring you have a seat at the table with your competition. Magazines, journals and newspapers are everywhere with unique readership and distribution channels. If programmed strategically, you'll increase your own audience base and have very noticeable ROI.
Of course print advertising is often not holistic enough for your modern marketing stack. Also, most print advertising packages also incentivize with many digital spots for exposure. Digital advertising can include web banners, email blasts to captive audiences, social media buys, geo-targeting and PPC ads. PPC, or pay-per-click, is just what it sounds like. Although it can be costly, you pay for every click from an ad directed toward your website. This is strong tool to gain fast results in a short time. Responsibly so, we believe with services like these you should start out at manageable levels and work your budget up to greater reach. Time is important when launching a brand, product or service, but we also cannot stress enough the need to do it right--never cutting corners and fulfilling the set marketing strategy every step of the way.
BrightBox believes advertising is best managed as a marketing tactic. It can be tailored uniquely for your business goals and marketing objectives. We leverage our media relationships to bring you the best return on your investment.


text: The bro Jo Jo Pellegrino is a F'N beast & always comes through with the bars! What do you think? Categories: Events, Music, PromotionBy The ConnectorSeptember 16, 2018Tags: barsbattle rapeminemmgkstaten islandvlad tv Author: The Connector https://richmondhood.com Post navigationPreviousPrevious post:Le Royale Tennis Mural by Jeremy Nieves, Mr.Tariq & Cody PrezNextNext post:Out Of Staten Island Interview w/ Mr.Tariq by Peanut The WickedRelated postsPersonal Protection Equipment – P.P.E. Packs Available Now!March 27, 2020Boat + Bridge 2/29/20 PackFebruary 19, 2020Robbin's Reef Lighthouse Visit w/ Noble Maritime Collection & Josh Pugliese,October 1, 2019Painkiller Paradise Video – Lou The HumanJuly 25, 2019Katiah – Tail Spin (Official Video)December 5, 2018Nectar SunglassesNovember 30, 2018


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Surrey Eagles Head Coach and Director of Player Personnel Brandon West has a unique perspective when it comes to leading a BCHL franchise.
Accepting a head coaching job in early December isn't easy, particularly when you have to move from one side of the country to the other. That's exactly what Jason Fortier did.
Kyle Wallach is currently enjoying a challenging season as an associate coach with the University of Vermont Men's Hockey Team. For Kyle and the rest of the staff at UVM, of course winning hockey games is important, but when you're overseeing a program that's built on the balance between academics and athletics, sometimes the bigger picture is far more crucial.
Kelowna Rockets Assistant Coach Kris Mallette has seen a lot in his career. From playing junior hockey with the Kelowna Rockets to coaching the Western Hockey League powerhouse, Kris played for nine teams in five leagues in under a decade.
Sven Butenschon is entering his second season as the head coach of the UBC Men's Hockey Team. Sven had tours of NHL duty with the Pittsburgh Penguins (more on that in a minute), the Edmonton Oilers, New York Islanders, and Vancouver Canucks before packing up his family to play a few more years in Europe.
Most Read:
1) JUNIOR B UPDATE: KIJHL's Castlegar Rebels announce new coach and GM; Sharp calling the shots for HJHL's Three Hills Thrashers
2) On Top of the World: CSSHL Keeps Gaining Traction in Canada's Hockey Landscape
3) Around the WHL: Eleven WHL players help Canada win Hlinka Gretzky gold; Tigers deal White to ICE
4) Meet Matthew Savoie, the NAX Forward Taking the CSSHL by Storm
5) Meet The Winners Of The 2018 HockeyNow Minor Hockey Player Of The Year Award Powered By Hockeyshot


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Indoor Wicker Coffee Table – Wicker furniture is definitely the most up-to-date trend in home furnishing about the globe. It is the hottest trend in Europe and America proper now, so much in order that celebrities' residences are sporting wicker furniture. It is the best thing ever because the level of innovation and talent which you can display with wicker is unparalleled. This is the reason designers are picking out wicker more than wood. Another explanation to obtain wicker furniture is definitely the eco-friendly nature of wicker. Wicker is just not wood and it isn't as destructive to the environment as wood is. Wicker is cultivated for the single purpose of making furniture out of it. This post related with 3 tier wicker basket stand.
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Arrivederci — Unisex T-Shirt
See you later! Those are the words we love to say when we are off to explore our next destination.
Soft and lightweight, with the right amount of stretch. It's comfortable and flattering for both men and women.
• 100% combed and ring-spun cotton (heather colors contain polyester)
• Fabric weight: 4.2 oz (142 g/m2)
• Shoulder-to-shoulder taping
• Side-seamed
Size Guide


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About Us
Who we are:
SCSM is a fitness apparel and accessories brand & online retailer. Our office is located in Charlotte, North Carolina in the heart of the city. With the support of our growing loyal customers we are able to impact-fully inspire others to live a lifestyle guided by fitness and faith.
Why we exist:
SCSM exists to share the message of hope to the hopeful but most importantly the hopeless. Over 45% of our country's population is physically inactive. If half of the parents in our country fall within this range the children will certainly be impacted.
Jeremiah 29:11
"For I know the thoughts that I think toward you, says the LORD, thoughts of peace and not of evil, to give you a future and a hope."
Other Causes:
Central to SCSM's mission to rep hope is our passion for children. Over 16 million American children don't get the food that they need on a regular basis. SCSM is committed to donating 10% of our net profits to combat child hunger in America and other causes. To put it simply we make quality clothes to help supply quality meals.
Our passion for extending hope to others remains. The creation of our clothing and media is not siply for the sake of success. Through the integrity of our brand we show people just how much we care by sharing messages of faith and representing hope. Join us in sharing the SCSM message.


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Red Currant is a New Zealand White doe who came to us in July 2018 as part of a big group of rabbits surrendered by a breeder who had been convinced to stop producing them for meat. We took in 9 of the 38 rabbits who were granted a stay of execution by being handed over to rescuers.
Red Currant is probably pregnant. I am fairly certain that I can feel babies in there and she has displayed an inclination towards nest building and appears to be eating for more than just herself. Time will tell. She will have to stay in the barn until we are certain one way or the other. As rabbits are only pregnant for thirty days it will pass quickly, and as I can already feel something it is likely to be well before the end of August.
In fact I actually thought she was going to have them the night she arrived as she was hopping around with a mouthful of hay trying to decide where to build a nest. I don't think she had ever had a choice in the matter before but in her new big hutch she has a number of cosy corners to choose from. We really hope I am wrong about her being pregnant because we have enough rabbits already but if she is at least her babies will be safe this time, and she can keep the females as her lifelong companions.


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LAHORE: Pakistan Muslim League-Quaid (PML-Q) leader Chaudhry Shujaat Hussain appeared before the National Accountability Bureau (NAB) on Thursday.
However, Chaudhry Pervaiz Elahi, ex-chief minister of Punjab, decided against appearing before the NAB officials.
The anti-graft body summoned the Chaudhry cousins yesterday in connection to allegations of owning assets beyond their known sources of income.
According to NAB sources, the Chaudhry cousins have failed to submit verified details of their assets. "If they do not submit the required records, NAB would initiate an inquiry against them under Schedule 2," the anti-corruption watchdog warned.
The Chaudhry cousins had earlier recorded their statements in NAB during previous hearings.
The duo are facing the charges of misusing their authorities, framed against them back in 2000 before they joined hands with former military ruler General Pervez Musharaf.
The graft case against Chaudhrys of Gujrat is also among the 179 mega corruption cases list, compiled and submitted in the Supreme Court by the anti-corruption watchdog in 2015.
The brothers were involved in Rs.2.428 billion corruption case, as per the details.
According to sources in NAB Lahore office, a number of shares have surfaced in various companies under the name of Pakistan Muslim League-Quaid leaders Chaudhry Shujaat Hussain, his cousin Chaudhry Pervez Elahi and their family.
Chaudhry Pervaiz Elahi is PTI's nominee for the post of Punjab Assembly speaker. He is contesting against Pakistan Muslim League-Nawaz's candidate Chaudhry Muhammad Iqbal Gujjar.
Chaudhry Shujaat Hussain appears before NAB in assets case
LAHORE: Pakistan Muslim League-Quaid (PML-Q) leader Chaudhry Shujaat Hussain appeared before the National Accountability Bureau (NAB) on Thursday.


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Another busy week in the Unicameral Legislature ended along with the month of January this past Friday. After a full week of hearings in the Agriculture, Business and Labor, and Health and Human Services Committees, extensive floor debate and many lunch or evening events, I capped off the week by speaking at the Blair Area Chamber of Commerce Annual Meeting. Senators receive a tremendous amount of information from a wide range of resources hoping to educate us about important issues. We hear from constituents like you reading this column and other individuals from around the state. We also are invited to events nearly every day of the week facilitated by organizations looking to spread the message of the work they do throughout the state.
This week I or my staff attended informational sessions sponsored by the National Guard Association, Habitat for Humanity, the Nebraska Chamber of Commerce, Voices for Children, the Nebraska Bankers, Nebraska Cattleman, American Bikers Aiming Towards Education, and the Blair Area Chamber of Commerce. These informational sessions are important functions of our roles as State Senators as we seek to be as informed as possible about how our decisions will affect all of Nebraska. Though I've developed specialties in the committees I'm a part of (listed above) I do appreciate learning more about other areas of policy. I especially appreciate hearing directly from constituents in Legislative District 16 when they come to Lincoln on their own, as a part of some of these organizations, or by contacting our office through phone or email.
Two of my bills, LB 312 and LB 381 came up for debate last week. Both bills were carried over after being introduced in the last session. LB 312 would expand opportunities for licensed dental hygienists, specifically in rural areas. We have a shortage of hygienists in rural Nebraska and my hope is this bill will expand access to care throughout the state. The bill was placed on Select File where we will discuss it once more before moving it on to Final Reading.
LB 381 would allow the state to move to a per-diem reimbursement process for state employees. It's estimated this bill would save the state substantial amounts of time and money by streamlining efficiencies in process and allowing state employees to spend time on more impactful work duties. This bill was placed on Final Reading where it will be voted on one more time before being sent to the Governor for his signature.
We continue to move through bills in committee as efficiently as possible. I encourage you to come testify at a public hearing for any bill you think is of particular interest or importance to you. This 60-day session is going fast and senators have been encouraged to make decisions on choosing a priority bill for the session. I've narrowed down my potential choices and hope to make a decision soon.
As always you can contact me at email@example.com or contact the office to speak with my Administrative Assistant, Ellie Stangl, or my Legislative Aide, Jacob Campbell. To read all introduced legislation please visit nebraskalegislature.gov. You may watch the live stream of the session when available at netnebraska.org.


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USU Resources
USU Policy 588: Resesarch Data This policy describes the rights and responsibilities of individual researchers and the institution in the use, retention, maintenance and sharing of data produced as part of USU's research enterprise.
USU Policy 587: Intellectual Property
Many sessions are offered at USU to help faculty and students develop data management skills. Below are links to past training sessions and/or the presentation files.
- Datapalooza! This half day event provides an overview of data management issues.
- Graduate Training Series, Oct. 17, 2016: How should I manage and store data effectively?
Download Presenter Slides
- Training for Research Faculty Series, Oct. 12, 2015: Open access to data: What do I need to know?
Download Presenter Slides; Download Workshop Audio
- Research Scholars Forum, Nov. 3,2016: Simple and effective methods for managing and sharing effective scientific data, given by Jeffery S. Horsburgh, Civil and Environmental Engineering.
- Merrill-Cazier Library Data Management Presentation, Fall 2015, given to academic departments at fall meetings.
USU People
Many people at USU are involved with data management. Finding the resources to help you with your data can be a challenge.
The Office of Research and Graduate Studies' Research Development Division has a wealth of information about funding, grant writing, workshops and seminars. The Proposal Development Specialists are familiar with the funding opportunities typically available in your college or school.
Feel free to email firstname.lastname@example.com if you need help finding information or are not sure to whom you should turn for information.
Other Resources
Information on Digitizing Herbarium Specimens, provided by Mary Barkworth. Multiple files are listed below:
- Creating Research Checklists
- Creating Teaching Checklists
- Image-vouchers Records
- PAk Field Records
- Personal Accounts in Symbiota Networks
- Preparing Herbarium Specimen Labels
- Preparing Images


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Most years, we don't travel for the Fourth of July. It's just easier to stay home, since all the places we would travel tend to be jam-packed with idiots.
This year, we spent the long weekend at home, just hanging out in the heat. Well, except for Saturday. Somehow I conned Hubby into letting me go see Eclipse. Actually, no conning was involved, I just had to tolerate some major eye rolling! I liked it, but it wasn't as good as the first two (same goes for the book) and I can't wait for Breaking Dawn, which was the best of the books and will hopefully be the best of the movies! I hear it will be in 2 parts.
Logan and I made homemade lemonade. We squeezed a lot of lemons! It was tasty but I'm not sure it was worth all the effort, really, for what we got out of it. That's sad to say, isn't it?
Popsicles were consumed in mass quantity, sometimes two at a time.
Since it was so darn hot, we spent a lot of time in the back yard, playing in kiddie pools and the sprinkler.
We also took a dip in the neighbor's big pool and on Monday, went to the gym to the Splash Pad spray park. Oh my goodness did the boys have a blast! It was Jordan's first time there and he was just splashing and dancing and giggling and hugging other kids. Pure joy!
A little video of Jordan tearing up the Splash Pad
All real men drive their John Deeres naked, right?
Here's me, all Little House on the Prairie or whatever.
We went to an outdoor church service on Sunday (a nest of barn swallows, a renegade mouse running through the crowd, car alarms, wind and screaming babies made for a really interesting time) and then had plans to go to the parade in North Branch. The heat got the best of Jordan, though, and he became hysterical so I put him down for a nap and Hubby stayed home with him. Logan and I went alone. It was fun, but hot!
Hubby had braved 8 Mile (yep, the one from the Eminem movie) to get some fireworks to do in our back yard, so we rationed those out over the weekend. Most were pretty lame. A couple of them were cool. We just figured we couldn't keep the baby out late enough to go to real fireworks. That would have been a mess. So we did these ones for Logan and he thought they were cool.
"I'm the king of New York!" (AKA the Statue of Liberty!)
A backyard toad hopped along and got in on the action. Yes, I put him in the garden before we lit the fireworks!
On Monday evening, just before dinnertime, Hubby came home from the gym and said "Grave Digger (the Monster Jam monster truck, for those of you who don't know) is up at Advanced Auto Parts." Well, color me redneck! The boys and I hopped out of the kiddie pool, threw on some clothes and we buzzed up there, arriving just in time before they closed up, so we got to see the "Grave Digger Experience."
I'm not quite sure who had the bigger smile when sitting in the driver's seat, Hubby or Logan.
The race champion throne
It was a cool thing for Logan to see, even though he pretended he didn't think it was cool, because he is really into Monster Jam. (By the way, what is UP with this whole "I can't show emotion when I'm excited" thing? It's SO annoying!!!!) He had to come home and play with his Grave Digger toy.
So, yeah, aside from the heat, which rendered the baby quite miserable and all of us but Logan unable to sleep at night, it was a very nice staycation with my guys!
Tuesday, July 6, 2010
4th of July Staycation
Most years, we don't travel for the Fourth of July. It's just easier to stay home, since all the places we would travel tend to be jam-packed with idiots.
Posted by Krystal at 6:50 AM
1 comments:
- Tina said...
I love being able to stay home and find fun things to do as a family. It looks like you all had a great time.
I'm going to see Eclipse tonight. I can't wait for it. I really wish she would write more Twilight books. The ending of Breaking Dawn was left a little open to where she could and you would think she would want that money.
- July 6, 2010 at 11:23 AM


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If you see the results of the Russian Trampoline Cup in Ramenskoje last week you will see two names on the winners list: Nikita Fedorenko and Irina Kundius.
Also you will recognize that Dimitri Ushakov is back in shape after missing the first Worldcup of the season.
More thinking about the names will bring you the a geographical question; from which city they all come from? Yeysk! Oh! Lets find it on Google Earth. Picturesque location on the shores of the Black Sea.
Done and add the knowledge about the head coaches: father and son Zaporozchenko.
They told us that the training is going better and better and so it is natural that they set high goals for Minsk.
Both Oleg Zaporozchenko good luck in Minsk and for the future…