BackgroundPeriodic stimulation of occipital areas using transcranial alternating current stimulation (tACS) at alpha (α) frequency (8–12 Hz) enhances electroencephalographic (EEG) α-oscillation long after tACS-offset. Two mechanisms have been suggested to underlie these changes in oscillatory EEG activity: tACS-induced entrainment of brain oscillations and/or tACS-induced changes in oscillatory circuits by spike-timing dependent plasticity.ObjectiveWe tested to what extent plasticity can account for tACS-aftereffects when controlling for entrainment “echoes.” To this end, we used a novel, intermittent tACS protocol and investigated the strength of the aftereffect as a function of phase continuity between successive tACS episodes, as well as the match between stimulation frequency and endogenous α-frequency.Methods12 healthy participants were stimulated at around individual α-frequency for 11–15 min in four sessions using intermittent tACS or sham. Successive tACS events were either phase-continuous or phase-discontinuous, and either 3 or 8 s long. EEG α-phase and power changes were compared after and between episodes of α-tACS across conditions and against sham.Resultsα-aftereffects were successfully replicated after intermittent stimulation using 8-s but not 3-s trains. These aftereffects did not reveal any of the characteristics of entrainment echoes in that they were independent of tACS phase-continuity and showed neither prolonged phase alignment nor frequency synchronization to the exact stimulation frequency.ConclusionOur results indicate that plasticity mechanisms are sufficient to explain α-aftereffects in response to α-tACS, and inform models of tACS-induced plasticity in oscillatory circuits. Modifying brain oscillations with tACS holds promise for clinical applications in disorders involving abnormal neural synchrony.
A number of rhythmic protocols have emerged for non-invasive brain stimulation (NIBS) in humans, including transcranial alternating current stimulation (tACS), oscillatory transcranial direct current stimulation (otDCS), and repetitive (also called rhythmic) transcranial magnetic stimulation (rTMS). With these techniques, it is possible to match the frequency of the externally applied electromagnetic fields to the intrinsic frequency of oscillatory neural population activity (“frequency-tuning”). Mounting evidence suggests that by this means tACS, otDCS, and rTMS can entrain brain oscillations and promote associated functions in a frequency-specific manner, in particular during (i.e., online to) stimulation. Here, we focus instead on the changes in oscillatory brain activity that persist after the end of stimulation. Understanding such aftereffects in healthy participants is an important step for developing these techniques into potentially useful clinical tools for the treatment of specific patient groups. Reviewing the electrophysiological evidence in healthy participants, we find aftereffects on brain oscillations to be a common outcome following tACS/otDCS and rTMS. However, we did not find a consistent, predictable pattern of aftereffects across studies, which is in contrast to the relative homogeneity of reported online effects. This indicates that aftereffects are partially dissociated from online, frequency-specific (entrainment) effects during tACS/otDCS and rTMS. We outline possible accounts and future directions for a better understanding of the link between online entrainment and offline aftereffects, which will be key for developing more targeted interventions into oscillatory brain activity.
BackgroundNavigation based on chemosensory information is one of the most important skills in the animal kingdom. Studies on odor localization suggest that humans have lost this ability. However, the experimental approaches used so far were limited to explicit judgements, which might ignore a residual ability for directional smelling on an implicit level without conscious appraisal.MethodsA novel cueing paradigm was developed in order to determine whether an implicit ability for directional smelling exists. Participants performed a visual two-alternative forced choice task in which the target was preceded either by a side-congruent or a side-incongruent olfactory spatial cue. An explicit odor localization task was implemented in a second experiment.ResultsNo effect of cue congruency on mean reaction times could be found. However, a time by condition interaction emerged, with significantly slower responses to congruently compared to incongruently cued targets at the beginning of the experiment. This cueing effect gradually disappeared throughout the course of the experiment. In addition, participants performed at chance level in the explicit odor localization task, thus confirming the results of previous research.ConclusionThe implicit cueing task suggests the existence of spatial information processing in the olfactory system. Response slowing after a side-congruent olfactory cue is interpreted as a cross-modal attentional interference effect. In addition, habituation might have led to a gradual disappearance of the cueing effect. It is concluded that under immobile conditions with passive monorhinal stimulation, humans are unable to explicitly determine the location of a pure odorant. Implicitly, however, odor localization seems to exert an influence on human behaviour. To our knowledge, these data are the first to show implicit effects of odor localization on overt human behaviour and thus support the hypothesis of residual directional smelling in humans.
Visuospatial attentional orienting has typically been studied in abstract tasks with low ecological validity. However, real-life tasks such as driving require allocation of working memory (WM) resources to several subtasks over and above orienting in a complex sensory environment. The aims of this study were twofold: firstly, to establish whether electrophysiological signatures of attentional orienting commonly observed under simplified task conditions generalize to a more naturalistic task situation with realistic-looking stimuli, and, secondly, to assess how these signatures are affected by increased WM load under such conditions. Sixteen healthy participants performed a dual task consisting of a spatial cueing paradigm and a concurrent verbal memory task that simulated aspects of an actual traffic situation. Behaviorally, we observed a load-induced detriment of sensitivity to targets. In the EEG, we replicated orienting-related alpha lateralization, the lateralized ERPs ADAN, EDAN, and LDAP, and the P1-N1 attention effect. When WM load was high (i.e., WM resources were reduced), lateralization of oscillatory activity in the lower alpha band was delayed. In the ERPs, we found that ADAN was also delayed, while EDAN was absent. Later ERP correlates were unaffected by load. Our results show that the findings in highly controlled artificial tasks can be generalized to spatial orienting in ecologically more valid tasks, and further suggest that the initiation of spatial orienting is delayed when WM demands of an unrelated secondary task are high.
Intersection accidents result in a significant proportion of road fatalities, and attention allocation likely plays a role. Attention allocation may depend on (limited) working memory (WM) capacity. Driving is often combined with tasks increasing WM load, consequently impairing attention orienting. This study (n = 22) investigated WM load effects on eventrelated potentials (ERPs) related to attention orienting. A simulated driving environment allowed continuous lane-keeping measurement. Participants were asked to orient attention covertly towards the side indicated by an arrow, and to respond only to moving cars appearing on the attended side by pressing a button. WM load was manipulated using a concurrent memory task. ERPs showed typical attentional modulation (cue: contralateral negativity, LDAP; car: N1, P1, SN, and P3) under low and high load conditions. With increased WM load, lane-keeping performance improved, while dual task performance degraded (memory task: increased error-rate; orienting task: increased false alarms, smaller P3).
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