Introducing the Prototypical Stimulus Characteristics Toolbox: Protosc.

Many studies use different categories of images to define their conditions. Since any difference between these categories is a valid candidate to explain category-related behavioral differences, knowledge about the objective image differences between categories is crucial for the interpretation of the behaviors. However, natural images vary in many image features and not every feature is equally important in describing the differences between the categories. Here, we provide a methodological approach to find as many of the image features as possible, using machine learning performance as a tool, that have predictive value over the category the images belong to. In other words, we describe a means to find the features of a group of images by which the categories can be objectively and quantitatively defined. Note that we are not aiming to provide a means for the best possible decoding performance; instead, our aim is to uncover prototypical characteristics of the categories. To facilitate the use of this method, we offer an open-source, MATLAB-based toolbox that performs such an analysis and aids the user in visualizing the features of relevance. We first applied the toolbox to a mock data set with a ground truth to show the sensitivity of the approach. Next, we applied the toolbox to a set of natural images as a more practical example.

Motor congruency and multisensory integration jointly facilitate visual information processing before movement execution.

Attention allows us to select important sensory information and enhances sensory information processing. Attention and our motor system are tightly coupled: attention is shifted to the target location before a goal-directed eye- or hand movement is executed. Congruent eye-hand movements to the same target can boost the effect of this pre-movement shift of attention. Moreover, visual information processing can be enhanced by, for example, auditory input presented in spatial and temporal proximity of visual input via multisensory integration (MSI). In this study, we investigated whether the combination of MSI and motor congruency can synergistically enhance visual information processing beyond what can be observed using motor congruency alone. Participants performed congruent eye- and hand movements during a 2-AFC visual discrimination task. The discrimination target was presented in the planning phase of the movements at the movement target location or a movement irrelevant location. Three conditions were compared: (1) a visual target without sound, (2) a visual target with sound spatially and temporally aligned (MSI) and (3) a visual target with sound temporally misaligned (no MSI). Performance was enhanced at the movement-relevant location when congruent motor actions and MSI coincide compared to the other conditions. Congruence in the motor system and MSI together therefore lead to enhanced sensory information processing beyond the effects of motor congruency alone, before a movement is executed. Such a synergy implies that the boost of attention previously observed for the independent factors is not at ceiling level, but can be increased even further when the right conditions are met.

Correction to: The disengagement of visual attention in the gap paradigm across adolescence.

In the manuscript, we write that saccadic reaction time was determined as the time between the offset of the last fixation on the central stimulus and the onset of the peripheral stimulus (the target).

Body representation does not lag behind in updating for the pubertal growth spurt.

Both making perceptual judgments about your own body and successfully moving your body through the world depend on a mental representation of the body. However, there are indications that moving might be challenging when your body is changing. For instance, the pubertal growth spurt has been reported to be negatively correlated to motor competence. A possible explanation for this clumsiness would be that when the body is growing fast, updating the body representation may lag behind, resulting in a mismatch between internal body representation and actual body size. The current study investigated this hypothesis by testing participants ranging from aged 6 to 50 years on both a tactile body image task and a motor body schema task. Separate groups of participants, including those in the age range when pubertal growth spurt occurs, were asked to estimate the distance between two simultaneously applied tactile stimuli on the arm and to move their hand through apertures of different widths. Tactile distance estimations were equal between participants before, during, and after the age range where the pubertal growth spurt is expected. Similarly, Bayesian evaluation of informative hypotheses showed that participants in the age range of the growth spurt did not move through the apertures as if their representation of the hand was smaller than its physical size. These results suggest that body representations do not lag behind in updating for the pubertal growth spurt.

The disengagement of visual attention in the gap paradigm across adolescence.

Attentional disengagement is important for successful interaction with our environment. The efficiency of attentional disengagement is commonly assessed using the gap paradigm. There is, however, a sharp contrast between the number of studies applying the gap paradigm to clinical populations and the knowledge about the underlying developmental trajectory of the gap effect. The aim of the present study was, therefore, to investigate attentional disengagement in a group of children aged 9-15. Besides the typically deployed gap and the overlap conditions, we also added a baseline condition in which the fixation point was removed at the moment that the target appeared. This allowed us to reveal the appropriate experimental conditions to unravel possible developmental differences. Correlational analyses showed that the size of the gap effect became smaller with increasing age, but only for the difference between the gap and the overlap conditions. This shows that there is a gradual increase in the capacity to disengage visual attention with increasing age, but that this effect only becomes apparent when the gap and the overlap conditions are compared. The gradual decrease of the gap effect with increasing age provides additional evidence that the attentional system becomes more efficient with increasing age and that this is a gradual process.

Visually Induced Inhibition of Return Affects the Integration of Auditory and Visual Information.

Multisensory integration (MSI) and exogenous spatial attention can both speedup responses to perceptual events. Recently, it has been shown that audiovisual integration at exogenously attended locations is reduced relative to unattended locations. This effect was observed at short cue-target intervals (200-250 ms). At longer intervals, however, the initial benefits of exogenous shifts of spatial attention at the cued location are often replaced by response time (RT) costs (also known as Inhibition of Return, IOR). Given these opposing cueing effects at shorter versus longer intervals, we decided to investigate whether MSI would also be affected by IOR. Uninformative exogenous visual spatial cues were presented between 350 and 450 ms prior to the onset of auditory, visual, and audiovisual targets. As expected, IOR was observed for visual targets (invalid cue RT < valid cue RT). For auditory and audiovisual targets, neither IOR nor any spatial cueing effects were observed. The amount of relative multisensory response enhancement and race model inequality violation was larger for uncued as compared with cued locations indicating that IOR reduces MSI. The results are discussed in the context of changes in unisensory signal strength at cued as compared with uncued locations.

Audiovisual integration in near and far space: effects of changes in distance and stimulus effectiveness.

A factor that is often not considered in multisensory research is the distance from which information is presented. Interestingly, various studies have shown that the distance at which information is presented can modulate the strength of multisensory interactions. In addition, our everyday multisensory experience in near and far space is rather asymmetrical in terms of retinal image size and stimulus intensity. This asymmetry is the result of the relation between the stimulus-observer distance and its retinal image size and intensity: an object that is further away is generally smaller on the retina as compared to the same object when it is presented nearer. Similarly, auditory intensity decreases as the distance from the observer increases. We investigated how each of these factors alone, and their combination, affected audiovisual integration. Unimodal and bimodal stimuli were presented in near and far space, with and without controlling for distance-dependent changes in retinal image size and intensity. Audiovisual integration was enhanced for stimuli that were presented in far space as compared to near space, but only when the stimuli were not corrected for visual angle and intensity. The same decrease in intensity and retinal size in near space did not enhance audiovisual integration, indicating that these results cannot be explained by changes in stimulus efficacy or an increase in distance alone, but rather by an interaction between these factors. The results are discussed in the context of multisensory experience and spatial uncertainty, and underline the importance of studying multisensory integration in the depth space.

Detection of lactate in the striatum without contamination of macromolecules by J-difference editing MRS at 7T.

Lactate levels are measurable by MRS and are related to neural activity. Therefore, it is of interest to accurately measure lactate levels in the basal ganglia networks. If sufficiently stable, lactate measurements may be used to investigate alterations in dopaminergic signalling in the striatum, facilitating the detection and diagnosis of metabolic deficits. The aim of this study is to provide a J-difference editing MRS technique for the selective editing of lactate only, thus allowing the detection of lactate without contamination of overlapping macromolecules. As a validation procedure, macromolecule nulling was combined with J-difference editing, and this was compared with J-difference editing with a new highly selective editing pulse. The use of a high-field (7T) MR scanner enables the application of editing pulses with very narrow bandwidth, which are selective for lactate. We show that, despite the sensitivity to B0 offsets, the use of a highly selective editing pulse is more efficient for the detection of lactate than the combination of a broad-band editing pulse with macromolecule nulling. Although the signal-to-noise ratio of uncontaminated lactate detection in healthy subjects is relatively low, this article describes the test-retest performance of lactate detection in the striatum when using highly selective J-difference editing MRS at 7 T. The coefficient of variation, σw and intraclass correlation coefficients for within- and between-subject differences of lactate were determined. Lactate levels in the left and right striatum were determined twice in 10 healthy volunteers. Despite the fact that the test-retest performance of lactate detection is moderate with a coefficient of variation of about 20% for lactate, these values can be used for the design of new studies comparing, for example, patient populations with healthy controls.

There is no attentional global effect: Attentional shifts are independent of the saccade endpoint.

Many studies have found a strong coupling between selective attention and eye movements. The premotor theory of attention suggests that saccade preparation is directly responsible for such attentional shifts. While it has already been shown that the attentional shift is not directly coupled to the final stages of motor execution, it is currently unknown to what aspect of the earlier stages of saccade preparation the attentional shift is coupled. An important step in this preparation process is resolving the landing point when multiple elements compete for the saccade. Here we ask how such a competition influences the presaccadic attentional locus and whether the presaccadic shift of attention is coupled to the saccade landing position or the possible saccade goals. To this end, we adopt a global effect paradigm where a target is accompanied by a salient distractor resulting in the majority of eye movements landing in between target and distractor. To determine the allocation of attention, participants are presented with a discrimination task shortly before the execution of the saccade. Despite a strong global effect obtained for saccade endpoints, we find little evidence for attentional facilitation at the location between target and distractor, but strong attentional facilitation at the location of the target and distractor. We argue that attention is coupled to active oculomotor programs, but not part of the resolution of these programs towards the execution of the saccade.

The influence of vertically and horizontally aligned visual distractors on aurally guided saccadic eye movements.

Eye movements towards a new target can be guided or disrupted by input from multiple modalities. The degree of oculomotor competition evoked by a distractor depends on both distractor and target properties, such as distractor salience or certainty regarding the target location. The ability to localize the target is particularly important when studying saccades made towards auditory targets, since determination of elevation and azimuth of a sound are based on different processes, and these processes may be affected independently by a distractor. We investigated the effects of a visual distractor on saccadic eye movements made to an auditory target in a two-dimensional plane. Results showed that the competition evoked by a vertical visual distractor was stronger compared with a horizontal visual distractor. The eye movements that were not captured by the vertical visual distractor were still influenced by it: a deviation of endpoints was seen in the direction of the visual distractor. Furthermore, the interference evoked by a high-contrast visual distractor was stronger compared with low-contrast visual stimuli, which was reflected by a faster initiation of an eye movement towards the high-contrast visual distractor and a stronger shift of endpoints in the direction of the high-contrast visual distractor. Together, these findings show that the influence of a visual distractor on aurally guided eye movements depends strongly on its location relative to the target, and to a lesser extent, on stimulus contrast.

The role of the frontal eye fields in oculomotor competition: image-guided TMS enhances contralateral target selection.

In order to execute a correct eye movement to a target in a search display, a saccade program toward the target element must be activated, while saccade programs toward distracting elements must be inhibited. The aim of the present study was to elucidate the role of the frontal eye fields (FEFs) in oculomotor competition. Functional magnetic resonance imaging-guided single-pulse transcranial magnetic stimulation (TMS) was administered over either the left FEF, the right FEF, or the vertex (control site) at 3 time intervals after target presentation, while subjects performed an oculomotor capture task. When TMS was applied over the FEF contralateral to the visual field where a target was presented, there was less interference of an ipsilateral distractor compared with FEF stimulation ipsilateral to the target's visual field or TMS over vertex. Furthermore, TMS over the FEFs decreased latencies of saccades to the contralateral visual field, irrespective of whether the saccade was directed to the target or to the distractor. These findings show that single-pulse TMS over the FEFs enhances the selection of a target in the contralateral visual field and decreases saccade latencies to the contralateral visual field.

Non-lateralized auditory input enhances averaged vectors in the oculomotor system.

The decision about which location should be the goal of the next eye movement is known to be determined by the interaction between auditory and visual input. This interaction can be explained by the vector theory that states that each element (either visual or auditory) in a scene evokes a vector in the oculomotor system. These vectors determine the direction in which the eye movement is initiated. Because auditory input is lateralized and localizable in most studies, it is currently unclear how non-lateralized auditory input interacts with the vectors evoked by visual input. In the current study, we investigated the influence of a non-lateralized auditory non-target on saccade accuracy (saccade angle deviation from the target) and latency in a single-target condition in Experiment 1 and a double-target condition in Experiment 2. The visual targets in Experiment 2 were positioned in such a way that saccades on average landed in between the two targets (i.e., a global effect). There was no effect of the auditory input on saccade accuracy in the single-target condition, but auditory input did influence saccade accuracy in the double-target condition. In both experiments, saccade latency increased when auditory input accompanied the visual target(s). Together, these findings show that non-lateralized auditory input enhances all vectors evoked by visual input. The results will be discussed in terms of their possible neural substrates.

A nasal visual field advantage in interocular competition.

When our eyes are confronted with discrepant images (yielding incompatible retinal inputs) interocular competition (IOC) is instigated. During IOC, one image temporarily dominates perception, while the other is suppressed. Many factors affecting IOC have been extensively examined. One factor that received surprisingly little attention, however, is the stimulus' visual hemifield (VHF) of origin. This is remarkable, as the VHF location of stimuli is known to affect visual performance in various contexts. Prompted by exploratory analyses, we examined five independent datasets of breaking continuous flash suppression experiments, to establish the VHF's role in IOC. We found that targets presented in nasal VHF locations broke through suppression much faster than targets in temporal VHF locations. Furthermore, we found that the magnitude of this nasal advantage depended on how strongly the targets were suppressed: the nasal advantage was larger for the recessive eye than for the dominant eye, and was larger in observers with a greater dominance imbalance between the eyes. Our findings suggest that the nasal advantage reported here originates in processing stages where IOC is resolved. Finally, we propose that a nasal advantage in IOC serves an adaptive role in human vision, as it can aid perception of partially occluded objects.

A global effect of capture saccades.

When two target elements are presented in close proximity, the endpoint of a saccade is generally positioned at an intermediate location ('global effect'). Here, we investigated whether the global effect also occurs for eye movements executed to distracting elements. To this end, we adapted the oculomotor capture paradigm such that on a subset of trials, two distractors were presented. When the two distractors were closely aligned, erroneous eye movements were initiated to a location in between the two distractors. Even though to a lesser extent, this effect was also present when the two distractors were presented further apart. In a second experiment, we investigated the global effect for eye movements in the presence of two targets. A strong global effect was observed when two targets were presented closely aligned, while this effect was absent when the targets were further apart. This study shows that there is a global effect when saccades are captured by distractors. This 'capture global' effect is different from the traditional global effect that occurs when two targets are presented because the global effect of capture saccades also occurs for remote elements. The spatial dynamics of this global effect will be explained in terms of the population coding theory.

The image features of emotional faces that predict the initial eye movement to a face.

Emotional facial expressions are important visual communication signals that indicate a sender's intent and emotional state to an observer. As such, it is not surprising that reactions to different expressions are thought to be automatic and independent of awareness. What is surprising, is that studies show inconsistent results concerning such automatic reactions, particularly when using different face stimuli. We argue that automatic reactions to facial expressions can be better explained, and better understood, in terms of quantitative descriptions of their low-level image features rather than in terms of the emotional content (e.g. angry) of the expressions. Here, we focused on overall spatial frequency (SF) and localized Histograms of Oriented Gradients (HOG) features. We used machine learning classification to reveal the SF and HOG features that are sufficient for classification of the initial eye movement towards one out of two simultaneously presented faces. Interestingly, the identified features serve as better predictors than the emotional content of the expressions. We therefore propose that our modelling approach can further specify which visual features drive these and other behavioural effects related to emotional expressions, which can help solve the inconsistencies found in this line of research.

Anomalous global effects induced by 'blind' distractors in visual hemifield defects.

Previous research has revealed that a stimulus presented in the blind visual field of participants with visual hemifield defects can evoke oculomotor competition, in the absence of awareness. Here we studied three cases to determine whether a distractor in a blind hemifield would be capable of inducing a global effect, a shift of saccade endpoint when target and distractor are close to each other, in participants with lesions of the optic radiations or striate cortex. We found that blind field distractors significantly shifted saccadic endpoints in two of three participants with lesions of either the striate cortex or distal optic radiations. The direction of the effect was paradoxical, however, in that saccadic endpoints shifted away from blind field distractors, whereas endpoints shifted towards distractors in the visible hemifields, which is the normal global effect. These results provide further evidence that elements presented in the blind visual field can generate modulatory interactions in the oculomotor system, which may differ from interactions in normal vision.

No direction specific costs in trans-saccadic memory.

Even though we frequently execute saccades, we perceive the external world as coherent and stable. An important mechanism of trans-saccadic perception is spatial remapping: the process of updating information across eye movements. Previous studies have indicated a right hemispheric dominance for spatial remapping, which has been proposed to translate into enhanced trans-saccadic memory for locations that are remapped into the right compared to the left hemisphere in healthy participants. Previous study designs suffered from several limitations, however (i.e. multiple eye movements had to be made instead of one, fixations were not controlled for, and ceiling effects were likely present). We therefore compared accuracy of trans-saccadic memory for central items after left- versus rightward eye movements, and secondary, for items that were remapped within the left versus right visual field. Participants memorized the location of a briefly presented item, made one saccade, and subsequently decided in what direction the item had shifted. We used a staircase to adjust task difficulty. Bayesian repeated measures ANOVAs were used to compare between left versus right eye movements and items in the left versus right visual field. We found most evidence against directional differences in trans-saccadic memory (BF10 = 0.23). We found some evidence suggestive of enhanced trans-saccadic memory for items that were remapped within the left compared to the right visual field (BF10 = 4.00). The latter result could be explained by a leftward spatial attention bias. As such, the hypothesized right hemispheric dominance for spatial remapping does not result in asymmetric trans-saccadic memory capacities in healthy participants.

Oculomotor capture in ADHD.

It is generally thought that deficits in response inhibition form an important area of dysfunction in patients with attention-deficit/hyperactivity disorder (ADHD). However, recent research using visual search paradigms seems to suggest that these inhibitory deficits do not extend towards inhibiting irrelevant distractors. Using an oculomotor capture task, the present study investigated whether boys with ADHD and their nonaffected brothers are impaired in suppressing reflexive eye movements to a task-irrelevant onset distractor. Results showed that boys with ADHD had slower responses than controls, but were as accurate in their eye movements as controls. Nonaffected brothers showed similar problems in the speed of responding as their affected brothers, which might suggest that this deficit relates to a familial risk for developing the disorder. Importantly, all three groups were equally captured by the distractor, which shows that boys with ADHD and their brothers are not more distracted by the distractor than are controls. Saccade latency and the proportion of intrusive saccades were related to continuous dimensions of ADHD symptoms, which suggests that these deficits are not simply present or absent, but rather indicate that the severity of these deficits relate to the severity of ADHD. The finding that boys with ADHD (and their nonaffected brothers) did not have problems inhibiting irrelevant distractors contradicts a general response inhibition deficiency in ADHD, which may be explained by the relatively independency of working memory in this type of response inhibition.

On the relative contributions of multisensory integration and crossmodal exogenous spatial attention to multisensory response enhancement.

Two processes that can give rise to multisensory response enhancement (MRE) are multisensory integration (MSI) and crossmodal exogenous spatial attention. It is, however, currently unclear what the relative contribution of each of these is to MRE. We investigated this issue using two tasks that are generally assumed to measure MSI (a redundant target effect task) and crossmodal exogenous spatial attention (a spatial cueing task). One block of trials consisted of unimodal auditory and visual targets designed to provide a unimodal baseline. In two other blocks of trials, the participants were presented with spatially and temporally aligned and misaligned audiovisual (AV) targets (0, 50, 100, and 200ms SOA). In the integration block, the participants were instructed to respond to the onset of the first target stimulus that they detected (A or V). The instruction for the cueing block was to respond only to the onset of the visual targets. The targets could appear at one of three locations: left, center, and right. The participants were instructed to respond only to lateral targets. The results indicated that MRE was caused by MSI at 0ms SOA. At 50ms SOA, both crossmodal exogenous spatial attention and MSI contributed to the observed MRE, whereas the MRE observed at the 100 and 200ms SOAs was attributable to crossmodal exogenous spatial attention, alerting, and temporal preparation. These results therefore suggest that there may be a temporal window in which both MSI and exogenous crossmodal spatial attention can contribute to multisensory response enhancement.

Exogenous spatial attention decreases audiovisual integration.

Multisensory integration (MSI) and spatial attention are both mechanisms through which the processing of sensory information can be facilitated. Studies on the interaction between spatial attention and MSI have mainly focused on the interaction between endogenous spatial attention and MSI. Most of these studies have shown that endogenously attending a multisensory target enhances MSI. It is currently unclear, however, whether and how exogenous spatial attention and MSI interact. In the current study, we investigated the interaction between these two important bottom-up processes in two experiments. In Experiment 1 the target location was task-relevant, and in Experiment 2 the target location was task-irrelevant. Valid or invalid exogenous auditory cues were presented before the onset of unimodal auditory, unimodal visual, and audiovisual targets. We observed reliable cueing effects and multisensory response enhancement in both experiments. To examine whether audiovisual integration was influenced by exogenous spatial attention, the amount of race model violation was compared between exogenously attended and unattended targets. In both Experiment 1 and Experiment 2, a decrease in MSI was observed when audiovisual targets were exogenously attended, compared to when they were not. The interaction between exogenous attention and MSI was less pronounced in Experiment 2. Therefore, our results indicate that exogenous attention diminishes MSI when spatial orienting is relevant. The results are discussed in terms of models of multisensory integration and attention.