Feature-based Attentional Amplitude Modulations of the Steady-state Visual Evoked Potentials Reflect Blood Oxygen Level Dependent Changes in Feature-sensitive Visual Areas.

Recent EEG studies have investigated basic principles of feature-based attention by means of frequency-tagged random dot kinematograms in which different colors are simultaneously presented at different temporal frequencies to elicit steady-state visual evoked potentials (SSVEPs). These experiments consistently showed global facilitation of the to-be-attended random dot kinematogram-a basic principle of feature-based attention. SSVEP source estimation suggested that posterior visual cortex from V1 to area hMT+/V5 is broadly activated by frequency-tagged stimuli. What is presently unknown is whether the feature-based attentional facilitation of SSVEPs is a rather unspecific neural response including all visual areas that follow the "on/off," or whether SSVEP feature-based amplitude enhancements are driven by activity in visual areas most sensitive to a specific feature, such as V4v in the case of color. Here, we leverage multimodal SSVEP-fMRI recordings in human participants and a multidimensional feature-based attention paradigm to investigate this question. Attending to shape produced significantly greater SSVEP-BOLD covariation in primary visual cortex compared with color. SSVEP-BOLD covariation during color selection increased along the visual hierarchy, with greatest values in areas V3 and V4. Importantly, in area hMT+/V5, we found no differences between shape and color selection. Results suggest that SSVEP amplitude enhancements in feature-based attention is not an unspecific enhancement of neural activity in all visual areas following the "on/off." These findings open new avenues to investigating neural dynamics of competitive interactions in specific visual areas sensitive to a certain feature in a more economical way and better temporal resolution compared with fMRI.

Feasibility of Digital Memory Assessments in an Unsupervised and Remote Study Setting.

Sensitive and frequent digital remote memory assessments via mobile devices hold the promise to facilitate the detection of cognitive impairment and decline. However, in order to be successful at scale, cognitive tests need to be applicable in unsupervised settings and confounding factors need to be understood. This study explored the feasibility of completely unsupervised digital cognitive assessments using three novel memory tasks in a Citizen Science project across Germany. To that end, the study aimed to identify factors associated with stronger participant retention, to examine test-retest reliability and the extent of practice effects, as well as to investigate the influence of uncontrolled settings such as time of day, delay between sessions or screen size on memory performance. A total of 1,407 adults (aged 18-89) participated in the study for up to 12 weeks, completing weekly memory tasks in addition to short questionnaires regarding sleep duration, subjective cognitive complaints as well as cold symptoms. Participation across memory tasks was pseudorandomized such that individuals were assigned to one of three memory paradigms resulting in three otherwise identical sub-studies. One hundred thirty-eight participants contributed to two of the three paradigms. Critically, for each memory task 12 independent parallel test sets were used to minimize effects of repeated testing. First, we observed a mean participant retention time of 44 days, or 4 active test sessions, and 77.5% compliance to the study protocol in an unsupervised setting with no contact between participants and study personnel, payment or feedback. We identified subject-level factors that contributed to higher retention times. Second, we found minor practice effects associated with repeated cognitive testing, and reveal evidence for acceptable-to-good retest reliability of mobile testing. Third, we show that memory performance assessed through repeated digital assessments was strongly associated with age in all paradigms, and individuals with subjectively reported cognitive decline presented lower mnemonic discrimination accuracy compared to non-complaining participants. Finally, we identified design-related factors that need to be incorporated in future studies such as the time delay between test sessions. Our results demonstrate the feasibility of fully unsupervised digital remote memory assessments and identify critical factors to account for in future studies.

Reducing the tendency for chronometric counting in duration discrimination tasks.

Chronometric counting is a prevalent issue in the study of human time perception as it reduces the construct validity of tasks and can conceal existing timing deficits. Several methods have been proposed to prevent counting strategies, but the factors promoting those strategies in specific tasks are largely uninvestigated. Here, we modified a classical two-interval duration discrimination task in two aspects that could affect the tendency to apply counting strategies. We removed the pause between the two intervals and changed the task instructions: Participants decided whether a short event occurred in the first or in the second half of a reference duration. In Experiment 1, both classical and modified task versions were performed under timing conditions, in which participants were asked not to count, and counting conditions, in which counting was explicitly instructed. The task modifications led to (i) a general decrease in judgment precision, (ii) a shift of the point of subjective equality, and (iii) a counting-related increase in reaction times, suggesting enhanced cognitive effort of counting during the modified task version. Precision in the two task versions was not differently affected by instructed counting. Experiment 2 demonstrates that-in the absence of any counting-related instructions-participants are less likely to engage in spontaneous counting in the modified task version. These results enhance our understanding of the two-interval duration discrimination task and demonstrate that the modifications tested here-although they do not significantly reduce the effectiveness of instructed counting-can diminish the spontaneous tendency to adopt counting strategies.

Shifting Attention in Feature Space: Fast Facilitation of the To-Be-Attended Feature Is Followed by Slow Inhibition of the To-Be-Ignored Feature.

In an explorative study, we investigated the time course of attentional selection shifts in feature-based attention in early visual cortex by means of steady-state visual evoked potentials (SSVEPs). To this end, we presented four flickering random dot kinematograms with red/blue, horizontal/vertical bars, respectively. Given the oscillatory nature of SSVEPs, we were able to investigate neural temporal dynamics of facilitation and inhibition/suppression when participants shifted attention either within (i.e., color to color) or between feature dimensions (i.e., color to orientation). Extending a previous study of our laboratory [Müller, M. M., Trautmann, M., & Keitel, C. Early visual cortex dynamics during top-down modulated shifts of feature-selective attention. Journal of Cognitive Neuroscience, 28, 643-655, 2016] to a full factorial design, we replicated a critical finding of our previous study: Facilitation of color was quickest, regardless of the origin of the shift (from color or orientation). Furthermore, facilitation of the newly to-be-attended and inhibition/suppression of the then to-be-ignored feature is not a time-invariant process that occurs instantaneously, but a biphasic one with longer time delays between the two processes. Interestingly, inhibition/suppression of the to-be-ignored feature after the shifting cue had a much longer latency with between- compared to within-dimensional shifts (by about 130-150 msec). The exploratory nature of our study is reasoned by two limiting factors: (a) Identical to our precursor study, we found no attentional SSVEP amplitude time course modulation for orientation, and (b) the signal-to-noise ratio for single trials was too poor to allow for reliable statistical testing of the latencies that were obtained with running t tests of averaged data.

Computing distance information from landmarks and self-motion cues - Differential contributions of anterior-lateral vs. posterior-medial entorhinal cortex in humans.

Landmarks and path integration cues are two important sources of spatial information for navigation. For example, both can be used to compute positional information, which, in rodents, has been related to computations in the entorhinal cortex. In humans, however, if and how the entorhinal cortex supports landmark-based navigation and path integration is poorly understood. To address this important question, we developed a novel spatial navigation task in which participants learned a target location and judged relative positions of test locations in relation to the target. Landmarks and path integration cues were dissociated, and their reliability levels were manipulated. Using fMRI adaptation, we investigated whether spatial distances among the test locations were encoded in the BOLD responses, separately for landmarks and self-motion cues. The results showed that the anterior-lateral entorhinal cortex adapted to the distance between successively visited test locations when landmarks were used for localization, meaning that its activation decreased as the distance between the currently occupied location and the preceding location decreased. In contrast, the posterior-medial entorhinal cortex adapted to between-location distance when path integration cues were used for localization. In addition, along with the hippocampus and the precuneus, both entorhinal subregions showed stronger activation in correct trials than incorrect trials, regardless of cue type and reliability level. Together, these findings suggest that subdivisions of entorhinal cortex encode fine-grained spatial information for different spatial cues, which provides important insights into how the entorhinal cortex supports different modes of spatial navigation.

Memory Image Completion: Establishing a task to behaviorally assess pattern completion in humans.

For memory retrieval, pattern completion is a crucial process that restores memories from partial or degraded cues. Neurocognitive aging models suggest that the aged memory system is biased toward pattern completion, resulting in a behavioral preference for retrieval over encoding of memories. Here, we built on our previously developed behavioral recognition memory paradigm-the Memory Image Completion (MIC) task-a task to specifically target pattern completion. First, we used the original design with concurrent eye-tracking in order to rule out perceptual confounds that could interact with recognition performance. Second, we developed parallel versions of the task to accommodate test settings in clinical environments or longitudinal studies. The results show that older adults have a deficit in pattern completion ability with a concurrent bias toward pattern completion. Importantly, eye-tracking data during encoding could not account for age-related performance differences. At retrieval, spatial viewing patterns for both age groups were more driven by stimulus identity than by response choice, but compared to young adults, older adults' fixation patterns overlapped more between stimuli that they (wrongly) thought had the same identity. This supports the observation that older adults choose responses perceived as similar to a learned stimulus, indicating a bias toward pattern completion. Additionally, two shorter versions of the task yielded comparable results, and no general learning effects were observed for repeated testing. Together, we present evidence that the MIC is a reliable behavioral task that targets pattern completion, that is easily and repeatedly applicable, and that is made freely available online.

The Human Dentate Gyrus Plays a Necessary Role in Discriminating New Memories.

Our day-to-day experiences are often similar to one another, occurring in the same place at the same time of day, with common people and objects, and with a shared purpose. Humans have an episodic memory to represent unique, personal events that are rich in detail [1]. For this to occur, at least two basic neural mechanisms are required: one to orthogonalize or "separate" overlapping input patterns at encoding and another to reinstate or "complete" memories from partial cues at retrieval [2-6]. To what extent do these purported "pattern separation" and "pattern completion" mechanisms rely on distinct subfields of the hippocampus [6]? Computational models [4-6] and lesion and genetic studies in rodents [7-12] largely point to the dentate gyrus as responsible for pattern separation and the CA3 and CA1 subfields for pattern completion (but see [13-16]). In high-resolution fMRI studies of humans, behavioral discrimination and completion tasks designed to approximate pattern separation and pattern completion, respectively, elicit the predicted pattern of activity in the dentate gyrus and CA3/CA1 [17-21]. Likewise, impaired behavioral discrimination has been demonstrated in individuals with hippocampal lesions [22, 23], but the lesions most likely encompass other subfields. Examination of these processes in individuals with selective lesions to hippocampal subfields is needed to infer causation [19]. Here, we report the rare case of BL, a 54-year-old man with bilateral ischemic lesions to the hippocampus [24] primarily affecting the dentate gyrus. Studying BL provides the unique opportunity to directly evaluate theories of hippocampal function that assign the dentate gyrus a specific role in discriminating old from new memories.

Changes in pattern completion--a key mechanism to explain age-related recognition memory deficits?

Accurate memory retrieval from partial or degraded input requires the reactivation of memory traces, a hippocampal mechanism termed pattern completion. Age-related changes in hippocampal integrity have been hypothesized to shift the balance of memory processes in favor of the retrieval of already stored information (pattern completion), to the detriment of encoding new events (pattern separation). Using a novel behavioral paradigm, we investigated the impact of cognitive aging (1) on recognition performance across different levels of stimulus completeness, and (2) on potential response biases. Participants were required to identify previously learned scenes among new ones. Additionally, all stimuli were presented in gradually masked versions to alter stimulus completeness. Both young and older adults performed increasingly poorly as the scenes became less complete, and this decline in performance was more pronounced in elderly participants indicative of a pattern completion deficit. Intriguingly, when novel scenes were shown, only the older adults showed an increased tendency to identify these as familiar scenes. In line with theoretical models, we argue that this reflects an age-related bias towards pattern completion.

Disgust- and not fear-evoking images hold our attention.

Even though disgust and fear are both negative emotions, they are characterized by different physiology and action tendencies. The aim of this study was to examine whether fear- and disgust-evoking images would produce different attention bias effects, specifically those related to attention (dis)engagement. Participants were asked to identify a target which was briefly presented around a central image cue, which could either be disgusting, frightening, or neutral. The interval between cue onset and target presentation varied within blocks (200, 500, 800, 1100 ms), allowing us to investigate the time course of attention engagement. Accuracy was lower and reaction times were longer when targets quickly (200 ms) followed disgust-evoking images than when they followed neutral- or fear-evoking images. For the other, longer interval conditions no significant image effects were found. These results suggest that emotion-specific attention effects can be found at very early visual processing stages and that only disgust-evoking images, and not fear-evoking ones, keep hold of our attention for longer. We speculate that this increase in early attention allocation is related to the need to perform a more comprehensive risk-assessment of the disgust-evoking images. The outcomes underline not only the importance of examining the time course of emotion induced attention effects but also the need to look beyond the dimensions of valence and arousal.

Subjective reports of stimulus, response, and decision times in speeded tasks: how accurate are decision time reports?

Four experiments examined how accurately participants can report the times of their own decisions. Within an auditory reaction time (RT) task, participants reported the time at which (a) the tone was presented, (b) they decided on the response, or (c) the response key was pressed. Decision time reports were checked for plausibility against the actual RTs, and we compared the effects of experimental manipulations on these two measures to see whether the reported decision times showed appropriate effects. In addition, we estimated the amount of error associated with individual decision time reports by checking how often participants' decision time reports were implausibly early (i.e., before stimulus onset) or late (i.e., after response), and by using several quantitative models. Overall, the results suggest that decision time reports are not very accurate but they may be usable for some purposes.