A Pressure Injection System for Investigating the Neuropharmacology of Information Processing in Awake Behaving Macaque Monkey Cortex

The top-down modulation of feed-forward cortical information processing is functionally important for many cognitive processes, including the modulation ...

German Primate Center

Disparate sensory streams originating from a common underlying event share similar dynamics, and this plays an important part in multisensory integration. Here we investigate audiovisual binding by presenting continuously changing, temporally congruent and incongruent stimuli. Recorded EEG signals are used to quantify spectrotemporal and waveform locking of neural activity to stimulus dynamics. Spectrotemporal analysis reveals locking to visual stimulus dynamics in both a broad alpha and the beta band. The properties of these effects suggest they are a correlate of bottom-up processing in the visual system. Waveform locking reveals two cortically distinct processes that lock to visual stimulus dynamics with differing topographies and time lags relative to the stimuli. Most importantly, these are modulated in strength by the congruency of an accompanying auditory stream. In addition, the waveform locking found at occipital electrodes shows an increase over stimulus duration for visual and congruent audiovisual stimuli. Hence we argue that these effects reflect audiovisual interaction. We thus propose that spectrotemporal and waveform locking reflect different mechanisms involved in the processing of dynamic audiovisual stimuli.

Visual stimulus locking of EEG is modulated by temporal congruency of auditory stimuli.

Although attention in older adults is an active research area, feature-selective aspects have not yet been explicitly studied. Here we report the results of an exploratory study involving directed changes in feature-selective attention. The stimuli used were two random dot kinematograms (RDKs) of different colours, superimposed and centrally presented. A colour cue with random onset after the beginning of each trial instructed young and older subjects to attend to one of the RDKs and detect short intervals of coherent motion while ignoring analogous motion events in the non-cued RDK. Behavioural data show that older adults could detect motion, but discriminated target from distracter motion less reliably than young adults. The method of frequency tagging allowed us to separate the EEG responses to the attended and ignored stimuli and directly compare steady-state visual evoked potential (SSVEP) amplitudes elicited by each stimulus before and after cue onset. We found that younger adults show a clear attentional enhancement of SSVEP amplitude in the post-cue interval, while older adults' SSVEP responses to attended and ignored stimuli do not differ. Thus, in situations where attentional selection cannot be spatially resolved, older adults show a deficit in selection that is not shared by young adults.

Feature-selective attention: evidence for a decline in old age.

Attention filters behaviorally relevant stimuli from the constant stream of sensory information comprising our environment. Research into underlying neural mechanisms in humans suggests that visual attention biases mutual suppression between stimuli resulting from competition for limited processing resources. As a consequence, processing of an attended stimulus is facilitated. This account makes 2 assumptions: 1) An attended stimulus is released from mutual suppression with competing stimuli and 2) an attended stimulus experiences greater gain in the presence of competing stimuli than when it is presented alone. Here, we tested these assumptions by recording frequency-tagged potentials elicited in early visual cortex that index stimulus-specific processing. We contrasted the processing of a given stimulus when its location was attended or unattended and in the presence or the absence of a nearby competing stimulus. At variance with previous findings, competition similarly suppressed processing of attended and unattended stimuli. Moreover, the magnitude of attentional gain was comparable in the presence or the absence of competing stimuli. We conclude that visuospatial selective attention does not directly modulate mutual suppression between stimuli but instead acts as a signal gain, which biases processing toward attended stimuli independent of competition.

Independent effects of attentional gain control and competitive interactions on visual stimulus processing.

Flickering stimuli evoke an oscillatory brain response with the same frequency as the driving stimulus, the so-called steady-state visual evoked potential (SSVEP). SSVEPs are robust brain signals whose amplitudes are enhanced with attention and thus play a major role in the development and use of non-invasive Brain-Computer Interfaces (BCIs). We compared the modulation of SSVEP amplitudes when subjects directly gazed at a flickering array of static dots (overt attention) to when they covertly shifted attention to the dots keeping their eyes at central fixation. A discrimination task was performed at the attended location to ensure that subjects shifted attention as instructed. Horizontal eye movements (allowed in overt attention but to be avoided in covert attention) were monitored by the horizontal electrooculogram. Subjects' behavioural performance was significantly reduced in covert attention compared to overt attention. Correspondingly, attentional modulation of SSVEP amplitudes by overt attention was larger in magnitude than for covert attention. Overt attention also changed the topographical distribution of SSVEP amplitudes on the scalp. Stimuli elicited the largest amplitudes at central occipital electrodes when they were overtly attended and at contralateral parieto-occipital sites when they were covertly attended. Accordingly, source analysis revealed clear centrally located sources in early visual areas in overt attention, regardless of the attended visual hemifield. Taken together these results affirm that overt and covert attention have qualitatively and quantitatively different effects on SSVEP responses as well as on task performance. Moreover, our results suggest that navigating SSVEP-BCIs with overt attention is more reliable and highlight some of the challenges in developing BCIs for patients who have lost the ability to move their eyes.

Effects of overt and covert attention on the steady-state visual evoked potential.

A better understanding of age-related change in the attentional modulation of perceptual processing may help elucidate cognitive change. For example, increased cognitive interference due to inappropriate processing of irrelevant information has been suggested to contribute to cognitive decline. However, it is not yet clear whether interference effects observed at later stages, such as executive function or response selection, are caused by leaky attentional selection at early, sensory stages of processing. Here, we investigated attentional control of sensory selection by comparing younger and older adults' ability to sustain spatial selective attention to one of two centrally presented, overlapping rapid serial visual presentation (RSVP) letter sequences, one large and one small. These stimuli elicited separable steady-state visual evoked potentials (SSVEP), which provide an index of early visual processing for each stimulus separately and are known to be modulated by selective attention. The condition of most interest required participants to attend to the larger letters while ignoring the smaller letters, as these foveally presented irrelevant stimuli were expected to present the strongest interference. Although the rapid presentation rates made the task demanding, detection ability did not differ between young and older adults. Accordingly, attentional modulation of SSVEP amplitudes was found in both age groups. Neither the magnitude nor the cortical sources of these SSVEP attention effects differed between age groups. Our results thus suggest that in the current task, the effect of voluntary spatial attention on sustained sensory processing in early visual areas is maintained in healthy old age.

Keeping focused: sustained spatial selective visual attention is maintained in healthy old age.

Performing a task across the left and right visual hemifields results in better performance than in a within-hemifield version of the task, termed the different-hemifield advantage. Although recent studies used transient stimuli that were presented with long ISIs, here we used a continuous objective electrophysiological (EEG) measure of competitive interactions for attentional processing resources in early visual cortex, the steady-state visual evoked potential (SSVEP). We frequency-tagged locations in each visual quadrant and at central fixation by flickering light-emitting diodes (LEDs) at different frequencies to elicit distinguishable SSVEPs. Stimuli were presented for several seconds, and participants were cued to attend to two LEDs either in one (Within) or distributed across left and right visual hemifields (Across). In addition, we introduced two reference measures: one for suppressive interactions between the peripheral LEDs by using a task at fixation where attention was withdrawn from the periphery and another estimating the upper bound of SSVEP amplitude by cueing participants to attend to only one of the peripheral LEDs. We found significantly greater SSVEP amplitude modulations in Across compared with Within hemifield conditions. No differences were found between SSVEP amplitudes elicited by the peripheral LEDs when participants attended to the centrally located LEDs compared with when peripheral LEDs had to be ignored in Across and Within trials. Attending to only one LED elicited the same SSVEP amplitude as Across conditions. Although behavioral data displayed a more complex pattern, SSVEP amplitudes were well in line with the predictions of the different-hemifield advantage account during sustained visuospatial attention.

Competitive interactions of attentional resources in early visual cortex during sustained visuospatial attention within or between visual hemifields: evidence for the different-hemifield advantage.

Deficient selection against irrelevant information has been proposed to underlie age-related cognitive decline. We recently reported evidence for maintained early sensory selection when older and younger adults used spatial selective attention to perform a challenging task. Here we explored age-related differences when spatial selection is not possible and feature-selective attention must be deployed. We additionally compared the integrity of feedforward processing by exploiting the well established phenomenon of suppression of visual cortical responses attributable to interstimulus competition. Electroencephalogram was measured while older and younger human adults responded to brief occurrences of coherent motion in an attended stimulus composed of randomly moving, orientation-defined, flickering bars. Attention was directed to horizontal or vertical bars by a pretrial cue, after which two orthogonally oriented, overlapping stimuli or a single stimulus were presented. Horizontal and vertical bars flickered at different frequencies and thereby elicited separable steady-state visual-evoked potentials, which were used to examine the effect of feature-based selection and the competitive influence of a second stimulus on ongoing visual processing. Age differences were found in feature-selective attentional modulation of visual responses: older adults did not show consistent modulation of magnitude or phase. In contrast, the suppressive effect of a second stimulus was robust and comparable in magnitude across age groups, suggesting that bottom-up processing of the current stimuli is essentially unchanged in healthy old age. Thus, it seems that visual processing per se is unchanged, but top-down attentional control is compromised in older adults when space cannot be used to guide selection.

Cliodhna Quigley

Cognitive Neuroscience Laboratory

German Primate Center

A Pressure Injection System for Investigating the Neuropharmacology of Information Processing in Awake Behaving Macaque Monkey Cortex

Cliodhna Quigley

.css-o250i3{font-size:18px;font-family:Open Sans,sans-serif;line-height:21.6px;}Cognitive Neuroscience Laboratory

German Primate Center

A Pressure Injection System for Investigating the Neuropharmacology of Information Processing in Awake Behaving Macaque Monkey Cortex

Cognitive Neuroscience Laboratory