We explored the nature and time course of effects generated by spatially uninformative peripheral cues by measuring these effects with localization responses to peripheral onsets or central arrow targets. In Experiment 1, participants made saccadic eye movements to equiprobable peripheral and central targets. At short cue-target onset asynchronies (CTOAs), responses to cued peripheral stimuli suffered from slowed responding attributable to sensory adaptation while responses to central targets were transiently facilitated, presumably due to cue-elicited oculomotor activation. At the longest CTOA, saccadic responses to central and peripheral targets were indistinguishably delayed, suggesting a common, output/decision effect (inhibition of return; IOR). In Experiment 2, we tested the hypothesis that the generation of this output effect is dependent on the activation state of the oculomotor system by forbidding eye movements and requiring keypress responses to frequent peripheral targets, while probing oculomotor behavior with saccades to infrequent central arrow targets. As predicted, saccades to central arrow targets showed neither the early facilitation nor later inhibitory effects that were robust in Experiment 1. At the long CTOA, manual responses to cued peripheral targets showed the typical delayed responses usually attributed to IOR. We recommend that this late "inhibitory" cueing effect (ICE) be distinguished from IOR because it lacks the cause (oculomotor activation) and effect (response bias) attributed to IOR when it was named by Posner, Rafal, Choate, and Vaughan (1985).
With two cueing tasks, in the present study we examined output-based inhibitory cueing effects (ICEs) with manual responses to arrow targets following manual or saccadic responses to arrow cues. In all experiments, ICEs were observed when manual localization responses were required to both the cues and targets, but only when the cue-target onset asynchrony (CTOA) was 2,000 ms or longer. In contrast, when saccadic responses were made in response to the cues, ICEs were only observed with CTOAs of 2,000 ms or less-and only when an auditory cue-back signal was used. The present study also showed that the magnitude of ICEs following saccadic responses to arrow cues decreased with time, much like traditional inhibition-of-return effects. The magnitude of ICEs following manual responses to arrow cues, however, appeared later in time and had no sign of decreasing even 3 s after cue onset. These findings suggest that ICEs linked to skeletomotor activation do exist and that the ICEs evoked by oculomotor activation can carry over to the skeletomotor system.
Inhibition of return is characterized by delayed responses to previously attended locations when the interval between stimuli is long enough. The present study employed steady-state visual evoked potentials (SSVEPs) as a measure of attentional modulation to explore the nature and time course of input- and output-based inhibitory cueing mechanisms that each slow response times at previously stimulated locations under different experimental conditions. The neural effects of behavioral inhibition were examined by comparing post-cue SSVEPs between cued and uncued locations measured across two tasks that differed only in the response modality (saccadic or manual response to targets). Grand averages of SSVEP amplitudes for each condition showed a reduction in amplitude at cued locations in the window of 100-500 ms post-cue, revealing an early, short-term decrease in the responses of neurons that can be attributed to sensory adaptation, regardless of response modality. Because primary visual cortex has been found to be one of the major sources of SSVEP signals, the results suggest that the SSVEP modulations observed were caused by input-based inhibition that occurred in V1, or visual areas earlier than V1, as a consequence of reduced visual input activity at previously cued locations. No SSVEP modulations were observed in either response condition late in the cue-target interval, suggesting that neither late input- nor output-based IOR modulates SSVEPs. These findings provide further electrophysiological support for the theory of multiple mechanisms contributing to behavioral cueing effects.
The neural systems that afford our ability to evaluate rewards and punishments are impacted by a variety of external factors. Here, we demonstrate that increased cognitive load reduces the functional efficacy of a reward processing system within the human medial-frontal cortex. In our paradigm, two groups of participants used performance feedback to estimate the exact duration of one second while electroencephalographic (EEG) data was recorded. Prior to performing the time estimation task, both groups were instructed to keep their eyes still and avoid blinking in line with well established EEG protocol. However, during performance of the time-estimation task, one of the two groups was provided with trial-to-trial-feedback about their performance on the time-estimation task and their eye movements to induce a higher level of cognitive load relative to participants in the other group who were solely provided with feedback about the accuracy of their temporal estimates. In line with previous work, we found that the higher level of cognitive load reduced the amplitude of the feedback-related negativity, a component of the human event-related brain potential associated with reward evaluation within the medial-frontal cortex. Importantly, our results provide further support that increased cognitive load reduces the functional efficacy of a neural system associated with reward processing.
To examine the relationship between visual imagery and autobiographical memory, eye position and pupil size were recorded while participants first searched for memories and then reconstructed the retrieved memories (Experiment 1), or only searched for memories (Experiment 2). In Experiment 1, we observed that, although recollective experience was not associated with the number of fixations per minute, memories that took longer to retrieve were linked to increased pupil size. In Experiment 2, we observed that directly retrieved memories were recalled more quickly and were accompanied by smaller pupils than generatively retrieved memories. After correcting for response time, retrieval mode also produced an effect, showing that decreased pupil size is not simply due to directly retrieved memories being recalled more quickly. These findings provide compelling evidence that objective measures, such as pupil size, can be used alongside subjective measures, such as self-reports, to distinguish between directly retrieved and generatively retrieved memories.
Studies of endogenous and exogenous attentional orienting in spatial cueing paradigms have been used to investigate inhibition of return, a behavioral phenomenon characterized by delayed reaction time in response to recently attended locations. When eye movements are suppressed, attention is covertly oriented to central or peripheral stimuli. Overt orienting, on the other hand, requires explicit eye movements to the stimuli. The present study examined the time course of slowed reaction times to previously attended locations when distractors are introduced into overt and covert orienting tasks. In a series of experiments, manual responses were required to targets following central and peripheral cues at three different cue-target intervals, with and without activated oculomotor systems. The results demonstrate that, when eye movements are suppressed, behavioral inhibition is reduced or delayed in magnitude by the presence of a distractor relative to conditions without distractors. However, the time course of behavioral inhibition when eye movements are required remains similar with or without distractors.
Inhibition of return (IOR) refers to an increase in reaction times to targets that appeared at a previously cued location relative to an uncued location, often investigated using a spatial cueing paradigm. Despite numerous studies that have examined many aspects of IOR, the neurophysiological mechanisms underlying IOR are still in dispute. The objective of the current research is to investigate the plausible mechanisms by manipulating the cue and target types between central and peripheral stimuli in a traditional cue-target paradigm with saccadic responses to targets. In peripheral-cueing conditions, we observed inhibitory cueing effects across all cue-target onset asynchronies (CTOAs) with peripheral targets, but IOR was smaller and arose later with central targets. No inhibition was observed in central-cueing conditions at any CTOAs. Empirical data were simulated using a two-dimensional dynamic neural field model. Our results and simulations support previous work demonstrating that, at short CTOAs, behavioral inhibition is only observed with repeated stimulation-an effect of sensory adaptation. With longer CTOAs, IOR is observed regardless of target type when peripheral cueing is used. Our findings suggest that behaviorally exhibited inhibitory cueing effects can be attributed to multiple mechanisms, including both attenuation of visual stimulation and local inhibition in the superior colliculus.
There are thought to be two forms of inhibition of return (IOR) depending on whether the oculomotor system is activated or suppressed. When saccades are allowed, output-based IOR is generated, whereas input-based IOR arises when saccades are prohibited. In a series of 4 experiments, we mixed or blocked compatible and incompatible trials with saccadic or manual responses to investigate whether cueing effects would follow the same pattern as those observed with more traditional peripheral onsets and central arrows. In all experiments, an uninformative cue was displayed, followed by a cue-back stimulus that was either red or green, indicating whether a compatible or incompatible response was required. The results showed that IOR was indeed observed for compatible responses in all tasks, whereas IOR was eliminated for incompatible trials-but only with saccadic responses. These findings indicate that the dissociation between input- and output-based forms of IOR depends on more than just oculomotor activation, providing further support for the existence of an inhibitory cueing effect that is distinct to the manual response modality.
Event-related potentials (ERPs) are tiny electrical brain responses in the human electroencephalogram that are typically not detectable until they are isolated by a process of signal averaging. Owing to the extremely smallsize of ERP components (ranging from less than 1 μV to tens of μV), compared to background brain rhythms, statistical analyses of ERPs are predominantly carried out in groups of subjects. This limitation is a barrier to the translation of ERP-based neuroscience to applications such as medical diagnostics. We show here that support vector machines (SVMs) are a useful method to detect ERP components in individual subjects with a small set of electrodes and a small number of trials for a mismatch negativity (MMN) ERP component. Such a reduced experiment setup is important for clinical applications. One hundred healthy individuals were presented with an auditory pattern containing pattern-violating deviants to evoke the MMN. Two-class SVMs were then trained to classify averaged ERP waveforms in response to the standard tone (tones that match the pattern) and deviant tone stimuli (tones that violate the pattern). The influence of kernel type, number of epochs, electrode selection, and temporal window size in the averaged waveform were explored. When using all electrodes, averages of all available epochs, and a temporal window from 0 to 900-ms post-stimulus, a linear SVM achieved 94.5 % accuracy. Further analyses using SVMs trained with narrower, sliding temporal windows confirmed the sensitivity of the SVM to data in the latency range associated with the MMN.
Inhibition of return (IOR) operationalizes a behavioral phenomenon characterized by slower responding to cued, relative to uncued, targets. Two independent forms of IOR have been theorized: input-based IOR occurs when the oculomotor system is quiescent, while output-based IOR occurs when the oculomotor system is engaged. EEG studies forbidding eye movements have demonstrated that reductions of target-elicited P1 components are correlated with IOR magnitude, but when eye movements occur, P1 effects bear no relationship to behavior. We expand on this work by adapting the cueing paradigm and recording event-related potentials: IOR is caused by oculomotor responses to central arrows or peripheral onsets and measured by key presses to peripheral targets. Behavioral IOR is observed in both conditions, but P1 reductions are absent in the central arrow condition. By contrast, arrow and peripheral cues enhance Nd, especially over contralateral electrode sites.
Event-related potentials (ERPs) may provide a non-invasive index of brain function for a range of clinical applications. However, as a lab-based technique, ERPs are limited by technical challenges that prevent full integration into clinical settings.