Strength of response suppression to distracter stimuli determines attentional-filtering performance in primate prefrontal neurons

Neurons in the primate dorsolateral prefrontal cortex (dlPFC) filter attended [corrected] targets distinctly from distracters through their response rates. The extent to which this ability correlates with the organism's performance, and the neural processes underlying it, remain unclear. We trained monkeys to attend to a visual target that differed in rank along a color-ordinal scale from that of a distracter. The animals' performance at focusing attention on the target and filtering out the distracter improved as ordinal distance between the stimuli increased. Importantly, dlPFC neurons also improved their filtering performance with increasing ordinal target-distracter distance; they built up their response rate in anticipation of the target-distracter onset, and then units encoding target representations increased their firing rate by similar amounts, whereas units encoding distracter representations gradually suppressed their rates as the interstimulus ordinal distance increased. These results suggest that attentional-filtering performance in primates relies upon dlPFC neurons' ability to suppress distracter representations.     

Investigating saccade programming in the praying mantis Tenodera aridifolia using distracter interference paradigms

To investigate the saccadic system in the mantis, I applied distracter interference paradigms. These involved presenting the mantis with a fixation target and one or several distracters supposed to affect saccades towards the target. When a single target was presented, a medium-sized target located in its lower visual field elicited higher rates of saccade response. This preference for target size and position was also observed when a target and a distracter were presented simultaneously. That is, the mantis chose and fixated the target rather than a distracter that was much smaller or larger than the target, or was located above the target. Furthermore, the mantis' preference was not affected by increasing the number of distracters. However, the presence of the distracter decreased the occurrence rate of saccade and increased the response time to saccade. I conclude that distracter interference paradigms are an effective way of investigating the visual processing underlying saccade generation in the mantis. Possible mechanisms of saccade generation in the mantis are discussed.     

Subitization and attentional engagement by transient stimuli

A series of experiments investigated the visual selection of moving and static items during enumeration. Small numbers of visual targets can be enumerated with little increase in reaction time and error with set size, a process referred to as 'subitization'. The number of items that can be subitized' is typically between one and four and known as the subitization range. This study looked for evidence of subitizing of subsets of items presented on a computer display. Fast and accurate enumeration was found for random configurations of moving targets even when presented among static distracters. This was not the case for static targets presented among moving or transient distracters. RTs to these targets were longer and showed a steady increase in RT with target number, even in the subitization range. However, when static targets and moving distracters were presented foveally, fast enumeration/subitization of the static targets was again possible. This was not due to reduced inter-item spacing, since linear effects of the number of targets still emerged when stimuli were presented peripherally but the size-spacing ratio was matched to the foveal presentations. There was indication that instead performance reflected perceived differences in movement speed for stimuli presented in parafoveal and more peripheral retinal regions. In support of this, subitization of static items improved as the movement speed of the distracters increased. The data suggest that the processes supporting subitization are highly sensitive to dynamic stimuli and depend on the ease of segmentation between static and moving arrays.     

Exploring BOLD Changes during Spatial Attention in Non-Stimulated Visual Cortex

Blood oxygen level-dependent (BOLD) responses were measured in parts of primary visual cortex that represented unstimulated visual field regions at different distances from a stimulated central target location. The composition of the visual scene varied by the presence or absence of additional peripheral distracter stimuli. Bottom-up effects were assessed by comparing peripheral activity during central stimulation vs. no stimulation. Top-down effects were assessed by comparing active vs. passive conditions. In passive conditions subjects simply watched the central letter stimuli and in active conditions they had to report occurrence of pre-defined targets in a rapid serial letter stream. Onset of the central letter stream enhanced activity in V1 representations of the stimulated region. Within representations of the periphery activation decreased and finally turned into deactivation with increasing distance from the stimulated location. This pattern was most pronounced in the active conditions and during the presence of peripheral stimuli. Active search for a target did not lead to additional enhancement at areas representing the attentional focus but to a stronger deactivation in the vicinity. Suppressed neuronal activity was also found in the non distracter condition suggesting a top-down attention driven effect. Our observations suggest that BOLD signal decreases in primary visual cortex are modulated by bottom-up sensory-driven factors such as the presence of distracters in the visual field as well as by top-down attentional processes.     

Properties of V1 neurons tuned to conjunctions of visual features: application of the V1 saliency hypothesis to visual search behavior

From a computational theory of V1, we formulate an optimization problem to investigate neural properties in the primary visual cortex (V1) from human reaction times (RTs) in visual search. The theory is the V1 saliency hypothesis that the bottom-up saliency of any visual location is represented by the highest V1 response to it relative to the background responses. The neural properties probed are those associated with the less known V1 neurons tuned simultaneously or conjunctively in two feature dimensions. The visual search is to find a target bar unique in color (C), orientation (O), motion direction (M), or redundantly in combinations of these features (e.g., CO, MO, or CM) among uniform background bars. A feature singleton target is salient because its evoked V1 response largely escapes the iso-feature suppression on responses to the background bars. The responses of the conjunctively tuned cells are manifested in the shortening of the RT for a redundant feature target (e.g., a CO target) from that predicted by a race between the RTs for the two corresponding single feature targets (e.g., C and O targets). Our investigation enables the following testable predictions. Contextual suppression on the response of a CO-tuned or MO-tuned conjunctive cell is weaker when the contextual inputs differ from the direct inputs in both feature dimensions, rather than just one. Additionally, CO-tuned cells and MO-tuned cells are often more active than the single feature tuned cells in response to the redundant feature targets, and this occurs more frequently for the MO-tuned cells such that the MO-tuned cells are no less likely than either the M-tuned or O-tuned neurons to be the most responsive neuron to dictate saliency for an MO target.     

Menstrual cycle shifts in attentional bias for courtship language

The current study investigated whether women show an attentional bias toward courtship language during the fertile phase of their menstrual cycle. Thirty heterosexual women (17 naturally cycling, 13 using hormonal contraceptives) completed a dichotic listening task on both a high and low fertility day of their menstrual cycle. Participants were asked to verbally repeat (shadow) an emotionally neutral target passage played in one ear while either a neutral or courtship distracter was played in the other ear. Courtship distracters were flirtatious in content but not overtly sexual. Shadowing errors were coded as a measure of attentional bias toward the distracter. Saliva samples were taken to determine whether levels of estradiol, progesterone and/or testosterone correlated with task performance. As predicted, naturally cycling women made more shadowing errors when listening to a courtship distracter during the fertile phase of their cycle than during the nonfertile phase. This effect was moderated by relationship status, such that fertile, mated women showed an attentional bias for courtship language but fertile single women did not. However, because of small sample sizes in the analysis, this relationship should be viewed as preliminary. Hormonal analysis revealed that higher levels of salivary estradiol predicted greater attentional bias toward courtship language in naturally cycling women. These results suggest that women's attention is drawn to verbal courtship signals when they are fertile, and that this shift is linked to increased estradiol release during the periovulatory phase.     

Selective and divided attention in animals

This article reviews some of the research on attentional processes in animals. In the traditional approach to selective attention, it is proposed that in addition to specific response attachments, animals also learn something about the dimension along which the stimuli fall (e.g., hue, brightness, or line orientation). More recently, there has been an attempt to find animal analogs to methodologies originally applied to research with humans. One line of research has been directed to the question of whether animals can locate a target among distracters faster if they are prepared for the presentation of the target (search image and priming). In the study of search image, the target is typically a food item and the cue consists of previous trials on which the same target is presented. In research on priming effects, the cue is typically different from the target but is a good predictor of its occurrence. The study of preattentive processes shows that perceptually, certain stimuli stand out from distracters better than others, depending not only on characteristics of the target relative to the distracters, but also on relations among the distracters. Research on divided attention is examined with the goal of determining whether an animal can process two elements of a compound sample with the same efficiency as one. Taken together, the reviewed research indicates that animals are capable of centrally (not just peripherally) attending to selective aspects of a stimulus display.     

Set-size effects for spatial frequency change and discrimination in multiple targets

In visual search tasks with a near-threshold target amongst distracters, log detection thresholds rise in proportion to the log of the number of stimuli. Previous research has shown a very steep slope for this set-size effect where the target is a change in spatial frequency (SF) across an ISI, suggesting a low-level explanation for 'change blindness (Wright et al., 2000). Here, we analyse stimulus and task variables in order to determine the contributions of stimulus detection and attention processes. Stimuli consisted of two 150 ms frames each containing 1 to 4 Gabor targets, with an ISI of 250 ms. In a 2AFC detection task with uniform distracters, slopes of 0.23-0.52 were found, in line with visual search results. 2AFC SF discrimination tasks gave slopes of 0.68, 0.69 with homogeneous distracters and 0.76-0.96 with inhomogeneous distracters, consistent with averaging of stimuli within a frame. If the distracters were also made to change across ISI, averaging was impossible, and focal attention was required to solve the discrimination. This always gave set-size slopes > 1. It is concluded that, under conditions where a stimulus array can be analysed globally, change detection performance is limited by signal detection mechanisms, rather than limited capacity attention or memory mechanisms. However, where this is prevented, for example by changing more than one item, limitations due to attention or memory produce an even steeper set-size effect.     

Multifocal attention filters targets from distracters within and beyond primate MT neurons' receptive field boundaries

Visual attention has been classically described as a spotlight that enhances the processing of a behaviorally relevant object. However, in many situations, humans and animals must simultaneously attend to several relevant objects separated by distracters. To account for this ability, various models of attention have been proposed including splitting of the attentional spotlight into multiple foci, zooming of the spotlight over a region of space, and switching of the spotlight among objects. We investigated this controversial issue by recording neuronal activity in visual area MT of two macaques while they attended to two translating objects that circumvented a third distracter object located inside the neurons' receptive field. We found that when the attended objects passed through or nearby the receptive field, neuronal responses to the distracter were either decreased or remained unaltered. These results demonstrate that attention can split into multiple spotlights corresponding to relevant objects while filtering out interspersed distracters.     

Time sharing in rats: A peak-interval procedure with gaps and distracters

Four hypotheses (switch, instructional-ambiguity, memory decay, and time sharing) were evaluated in a reversed peak-interval procedure with gaps by presenting distracter stimuli during the uninterrupted timed signal. The switch, instructional-ambiguity, and memory-decay hypotheses predict that subjects should time through the distracter and delay responding during gaps. The time-sharing hypothesis assumes that the internal clock shares attentional and working-memory resources with other processes, so that both gaps and distracters delay timing by causing working memory to decay. We found that response functions were displaced both by gaps and by distracters. Computer simulations show that when combined, the memory-decay and time-sharing hypotheses can mechanistically address present data, suggesting that these two hypotheses may reflect different levels of analysis of the same phenomenon.     

The Presence of a Therapy Dog Results in Improved Object Recognition Performance in Preschool Children

ABSTRACT

The purpose of this study was to determine whether the presence of a dog would have an impact on object recognition memory performance of preschool children. This work represents an extension of previous research which found that preschoolers require fewer instructional prompts to complete this type of memory task when in the presence of a dog. If children require fewer instructional prompts it is possible that they are better able to focus on the task itself and as a result, improved memory performance is likely. Because the earlier experiment utilized a very simple version of the task that was readily completed by the preschool children, the overall performance data were at ceiling. The current study, involving 20 preschool children, included a manipulation of task difficulty through varying the number of distracters (one versus four) present at test. Increasing the number of distractors in a simple recognition task is known to make that task more challenging, and thus performance was expected to be slower and less accurate in the four distracter conditions relative to the one-distracter conditions. The collaborators in the study were either a therapy dog or a human. A two-way repeated measures design was used such that each child served as his/her own control and was tested in each of four separate conditions: dog present (one and four distracters) and human present (one and four distracters). The results showed that the preschool children performed the object recognition task faster and more accurately in the presence of the therapy dog relative to a human and also in the one-distracter versus four-distracter condition. The authors conclude that these effects result from increased focus and/or motivation resulting from the presence of the dog.     

Spatial interactions in simple and combined-feature visual search.

To isolate the mechanisms responsible for spatial interactions in visual search, we investigated the effects of inter-element distance and positional jittering in both simple (in/s) and combined-feature (inverted V in Vs) search tasks in which the observer had to find a target in a background of similar nontargets. Thresholds, defined as the stimulus duration for 75% correct, were measured for 'present' and 'absent' target conditions as a function of background numerosity (ranging from 4 to 64 background elements), independently for four inter-element distances and three positional jittering conditions. Results show: (1) both simple and combined-feature search involve a parallel, capacity limited process, (2) thresholds for parallel search of simple features are directly related to inter-element distance whereas this has little effect on thresholds in combined-feature search, and (3) positional jittering has a direct effect on thresholds in combined-feature search and an inverse effect in simple-feature search. These results indicate that two different mechanisms of spatial interactions are involved in parallel search. The activation of each of the two mechanisms depends on the stimulus used. In parallel search for simple and dissimilar features, the underlying mechanism is a short-range one, based on lateral inhibition, whereas the parallel search for combined and similar features is based on a nontarget grouping mechanism which relies on facilitatory interactions between regular elements.     

Reduced dNTP interaction of human immunodeficiency virus type 1 reverse transcriptase promotes strand transfer

We have recently demonstrated that HIV-1 RT mutants characterized by low dNTP binding affinity display significantly reduced dNTP incorporation kinetics in comparison to wild-type RT. This defect is particularly emphasized at low dNTP concentrations where WT RT remains capable of efficient synthesis. Kinetic interference in DNA synthesis can induce RT pausing and slow down the synthesis rate. RT stalling and slow synthesis rate can enhance RNA template cleavage by RT-RNase H, facilitating transfer of the primer to a homologous template. We therefore hypothesized that reduced dNTP binding RT mutants can promote template switching during minus strand synthesis more efficiently than WT HIV-1 RT at low dNTP concentrations. To test this hypothesis, we employed two dNTP binding HIV-1 RT mutants, Q151N and V148I. Indeed, as the dNTP concentration was decreased, the template switching frequency progressively increased for both WT and mutant RTs. However, as predicted, the RT mutants promoted more transfers compared with WT RT. The WT and mutant RTs were similar in their intrinsic RNase H activity, supporting that the elevated template switching efficiency of the mutants was not the result of the mutations enhancing RNase H activity. Rather, kinetic interference leading to stalled DNA synthesis likely enhanced transfers. These results suggest that the RT-dNTP substrate interaction mechanistically influences strand transfer and recombination of HIV-1 RT.     

Attractor models of working memory and their modulation by reward

This work reports an empirical examination of two key issues in theoretical neuroscience: distractibility in the context of working memory (WM) and its reward dependence. While these issues have been examined fruitfully in isolation (e.g. Macoveanu et al. in Biol Cybern 96(4): 407–19, 2007), we address them here in tandem, with a focus on how distractibility and reward interact. In particular, we parameterise an observation model that embodies the nonlinear form of such interactions, as described in a recent neuronal network model (Gruber et al. in J Comput Neurosci 20:153–166, 2006). We observe that memory for a target stimulus can be corrupted by distracters in the delay period. Interestingly, in contrast to our theoretical predictions, this corruption was only partial. Distracters do not simply overwrite target; rather, a compromise is reached between target and distracter. Finally, we observed a trend towards a reduced distractibility under conditions of high reward. We discuss the implications of these findings for theoretical formulations of basal and dopamine (DA)-modulated neural bump- attractor networks of working memory.     

Electrophysiological evidence for immature processing capacity and filtering in visuospatial working memory in adolescents

The present study investigated the development of visuospatial working memory (VSWM) capacity and the efficiency of filtering in VSWM in adolescence. To this end, a group of IQ-matched adults and adolescents performed a VSWM change detection task with manipulations of WM-load and distraction, while performance and electrophysiological contralateral delay activity (CDA) were measured. The CDA is a lateralized ERP marker of the number of targets and distracters that are selectively encoded/maintained in WM from one hemifield of the memory display. Significantly lower VSWM-capacity (Cowan's K) was found in adolescents than adults, and adolescents' WM performance (in terms of accuracy and speed) also suffered more from the presence of distracters. Distracter-related CDA responses were partly indicative of higher distracter encoding/maintenance in WM in adolescents and were positively correlated with performance measures of distracter interference. This correlation suggests that the higher interference of distracters on WM performance in adolescents was caused by an inability to block distracters from processing and maintenance in WM. The lower visuospatial WM-capacity (K) in adolescents in the high load (3 items) condition was accompanied by a trend (p<.10) towards higher CDA amplitudes in adolescents than adults, whereas CDA amplitudes in the low load (1 item) condition were comparable between adolescents and adults. These findings point to immaturity of frontal-parietal WM-attention networks that support visuospatial WM processing in adolescence.     

The Antisaccade Task: Visual Distractors Elicit a Location-Independent Planning ‘Cost’

The presentation of a remote – but not proximal – distractor concurrent with target onset increases prosaccade reaction times (RT) (i.e., the remote distractor effect: RDE). The competitive integration model asserts that the RDE represents the time required to resolve the conflict for a common saccade threshold between target- and distractor-related saccade generating commands in the superior colliculus. To our knowledge however, no previous research has examined whether remote and proximal distractors differentially influence antisaccade RTs. This represents a notable question because antisaccades require decoupling of the spatial relations between stimulus and response (SR) and therefore provide a basis for determining whether the sensory- and/or motor-related features of a distractor influence response planning. Participants completed pro- and antisaccades in a target-only condition and conditions wherein the target was concurrently presented with a proximal or remote distractor. As expected, prosaccade RTs elicited a reliable RDE. In contrast, antisaccade RTs were increased independent of the distractor’s spatial location and the magnitude of the effect was comparable across each distractor location. Thus, distractor-related antisaccade RT costs are not accounted for by a competitive integration between conflicting saccade generating commands. Instead, we propose that a visual distractor increases uncertainty related to the evocation of the response-selection rule necessary for decoupling SR relations.