Olfactory discrimination: when vision matters?

Many previous studies have attempted to investigate the effect of visual cues on olfactory perception in humans. The majority of this research has only looked at the modulatory effect of color, which has typically been explained in terms of multisensory perceptual interactions. However, such crossmodal effects may equally well relate to interactions taking place at a higher level of information processing as well. In fact, it is well-known that semantic knowledge can have a substantial effect on people's olfactory perception. In the present study, we therefore investigated the influence of visual cues, consisting of color patches and/or shapes, on people's olfactory discrimination performance. Participants had to make speeded odor discrimination responses (lemon vs. strawberry) while viewing a red or yellow color patch, an outline drawing of a strawberry or lemon, or a combination of these color and shape cues. Even though participants were instructed to ignore the visual stimuli, our results demonstrate that the accuracy of their odor discrimination responses was influenced by visual distractors. This result shows that both color and shape information are taken into account during speeded olfactory discrimination, even when such information is completely task irrelevant, hinting at the automaticity of such higher level visual-olfactory crossmodal interactions.     

Visual Experience Determines the Use of External Reference Frames in Joint Action Control

Vision plays a crucial role in human interaction by facilitating the coordination of one''s own actions with those of others in space and time. While previous findings have demonstrated that vision determines the default use of reference frames, little is known about the role of visual experience in coding action-space during joint action. Here, we tested if and how visual experience influences the use of reference frames in joint action control. Dyads of congenitally-blind, blindfolded-sighted, and seeing individuals took part in an auditory version of the social Simon task, which required each participant to respond to one of two sounds presented to the left or right of both participants. To disentangle the contribution of external—agent-based and response-based—reference frames during joint action, participants performed the task with their respective response (right) hands uncrossed or crossed over one another. Although the location of the auditory stimulus was completely task-irrelevant, participants responded overall faster when the stimulus location spatially corresponded to the required response side than when they were spatially non-corresponding: a phenomenon known as the social Simon effect (SSE). In sighted participants, the SSE occurred irrespective of whether hands were crossed or uncrossed, suggesting the use of external, response-based reference frames. Congenitally-blind participants also showed an SSE, but only with uncrossed hands. We argue that congenitally-blind people use both agent-based and response-based reference frames resulting in conflicting spatial information when hands are crossed and, thus, canceling out the SSE. These results imply that joint action control functions on the basis of external reference frames independent of the presence or (transient/permanent) absence of vision. However, the type of external reference frames used for organizing motor control in joint action seems to be determined by visual experience.     

The Contribution of Mere Recognition to the P300 Effect in a Concealed Information Test

In two experiments, we investigated the role of mere recognition in a P300 based CIT. Mere recognition was isolated by having participants respond based on an irrelevant dimension of the stimuli. In Experiment 1 stimuli consisted of familiar and unfamiliar faces, with a dot placed on the left or the right cheeck. Participants responded according to dot location. In the second experiment, participants were presented with autobiographical information, alternated with irrelevant stimuli, while instructed to respond based on the case of the stimuli. Results showed that with both familiar faces, and autobiographical information, mere recognition was sufficient to elicit a P300.

Functional Difference between Sustained and Transient Modulations of Cognitive Control in the Simon Task: Evidence from False Alarm Responses on No-Go Trials

Cognitive control in response compatibility tasks is modulated by the task context. Two types of contextual modulations have been demonstrated; sustained (block-wise) and transient (trial-by-trial). Recent research suggests that these modulations have different underlying mechanisms. This study presents new evidence supporting this claim by comparing false alarm (FA) responses on no-go trials of the Simon task between the sustained and transient contexts. In Experiment 1, the sustained context was manipulated so that a block included a larger number of incongruent trials. Results showed that participants made more FA responses by the hand opposite to the stimulus location. This suggests a generation of response bias in which the task-irrelevant location information is utilized in a reversed manner (i.e., to respond with the right hand to a stimulus presented on the left side and vice versa). Next, Experiment 2 examined the effect of the transient context and found that overall FA rate was lower when a no-go trial was preceded by an incongruent trial than by a congruent trial, whereas such response bias as that shown in Experiment 1 was not demonstrated. This suggests that the transient conflict context enhances inhibition of the task-irrelevant process but does not make the task-irrelevant information actively usable. Based on these results, we propound two types of cognitive control modulations as adaptive behaviors: response biasing based on utilization of the task-irrelevant information under the sustained conflict context and transient enhancement of inhibition of the task-irrelevant process based on the online conflict monitoring.     

Human implicit memory for irrelevant dimension values is similar to rats' incidental memory in simultaneous discrimination tasks

Participants completed a category-learning task in which they needed to discover which of three stimulus dimensions (shape, color or size) was relevant. After meeting a learning criterion (nine of 10 consecutive correct responses), participants continued making categorization choices and response latencies associated with these trials were examined. In both Experiments 1 and 2, people responded reliably faster when correct responses matched the previous responses with respect to irrelevant dimension values. Thus, they demonstrated a form of incidental short-term memory analogous to that we previously reported in studies of rats. In Experiment 2, participants' explicit memory for irrelevant dimension values was assessed after category learning was complete. The results indicated that people were unaware of the irrelevant dimension values encountered on trials preceding surprise probe trials. This indicates that memory for the irrelevant dimension values was implicit (i.e. unconscious). The findings are discussed with respect to both human and non-human studies of hippocampus-independent memory and implicit memory.     

Rat spatial memory tasks adapted for humans: characterization in subjects with intact brain and subjects with selective medial temporal lobe thermal lesions

In the present paper we describe five tests, 3 of which were designed to be similar to tasks used with rodents. Results obtained from control subjects, patients with selective thermo-coagulation lesions to the medial temporal lobe and results from non-human primates and rodents are discussed. The tests involve memory for spatial locations acquired by moving around in a room, memory for objects subjects interacted with, or memory for objects and their locations. Two of the spatial memory tasks were designed specifically as analogs of the Morris water task and the 8-arm radial-maze tasks used with rats. The Morris water task was modeled by hiding a sensor under the carpet of a room (Invisible Sensor Task). Subjects had to learn its location by using an array of visual cues available in the room. A path integration task was developed in order to study the non-visual acquisition of a cognitive representation of the spatial location of objects. In the non-visual spatial memory task, we blindfolded subjects and led them to a room where they had to find 3 objects and remember their locations. We designed an object location task by placing 4 objects in a room that subjects observed for later recall of their locations. A recognition task, and a novelty detection task were given subsequent to the recall task. An 8-arm radial-maze was recreated by placing stands at equal distance from each other around the room, and asking subjects to visit each stand once, from a central point. A non-spatial working memory task was designed to be the non-spatial equivalent of the radial maze. Search paths recorded on the first trial of the Invisible Sensor Task, when subjects search for the target by trial and error are reported. An analysis of the search paths revealed that patients with lesions to the right or left hippocampus or parahippocampal cortex employed the same type of search strategies as normal controls did, showing similarities and differences to the search behavior recorded in rats. Interestingly, patients with lesions that included the right parahippocampal cortex were impaired relative to patients with lesions to the right hippocampus that spared the parahippocampal cortex, when recall of the sensor was tested after a 30 min delay (Bohbot et al. 1998). No differences were obtained between control subjects and patients with selective thermal lesions to the medial temporal lobe, when tested on the radial-maze, the non-spatial analogue to the radial-maze and the path integration tasks. Differences in methodological procedures, learning strategies and lesion location could account for some of the discrepant results between humans and non-human species. Patients with lesions to the right hippocampus, irrespective of whether the right parahippocampal cortex was spared or damaged, had difficulties remembering the particular configuration and identity of objects in the novelty detection of the object location task. This supports the role of the human right hippocampus for spatial memory, in this case, involving memory for the location of elements in the room; learning known to require the hippocampus in the rat.     

Feature-Specific Encoding Flexibility in Visual Working Memory

The current study examined selective encoding in visual working memory by systematically investigating interference from task-irrelevant features. The stimuli were objects defined by three features (color, shape, and location), and during a delay period, any of the features could switch between two objects. Additionally, single- and whole-probe trials were randomized within experimental blocks to investigate effects of memory retrieval. A series of relevant-feature switch detection tasks, where one feature was task-irrelevant, showed that interference from the task-irrelevant feature was only observed in the color-shape task, suggesting that color and shape information could be successfully filtered out, but location information could not, even when location was a task-irrelevant feature. Therefore, although location information is added to object representations independent of task demands in a relatively automatic manner, other features (e.g., color, shape) can be flexibly added to object representations.     

Sustained Spatial Attention to Vibrotactile Stimulation in the Flutter Range: Relevant Brain Regions and Their Interaction

The present functional magnetic resonance imaging (fMRI) study was designed to get a better understanding of the brain regions involved in sustained spatial attention to tactile events and to ascertain to what extent their activation was correlated. We presented continuous 20 Hz vibrotactile stimuli (range of flutter) concurrently to the left and right index fingers of healthy human volunteers. An arrow cue instructed subjects in a trial-by-trial fashion to attend to the left or right index finger and to detect rare target events that were embedded in the vibrotactile stimulation streams. We found blood oxygen level-dependent (BOLD) attentional modulation in primary somatosensory cortex (SI), mainly covering Brodmann area 1, 2, and 3b, as well as in secondary somatosensory cortex (SII), contralateral to the to-be-attended hand. Furthermore, attention to the right (dominant) hand resulted in additional BOLD modulation in left posterior insula. All of the effects were caused by an increased activation when attention was paid to the contralateral hand, except for the effects in left SI and insula. In left SI, the effect was related to a mixture of both a slight increase in activation when attention was paid to the contralateral hand as well as a slight decrease in activation when attention was paid to the ipsilateral hand (i.e., the tactile distraction condition). In contrast, the effect in left posterior insula was exclusively driven by a relative decrease in activation in the tactile distraction condition, which points to an active inhibition when tactile information is irrelevant. Finally, correlation analyses indicate a linear relationship between attention effects in intrahemispheric somatosensory cortices, since attentional modulation in SI and SII were interrelated within one hemisphere but not across hemispheres. All in all, our results provide a basis for future research on sustained attention to continuous vibrotactile stimulation in the range of flutter.     

Cognitive Bias in Ambiguity Judgements: Using Computational Models to Dissect the Effects of Mild Mood Manipulation in Humans

Positive and negative moods can be treated as prior expectations over future delivery of rewards and punishments. This provides an inferential foundation for the cognitive (judgement) bias task, now widely-used for assessing affective states in non-human animals. In the task, information about affect is extracted from the optimistic or pessimistic manner in which participants resolve ambiguities in sensory input. Here, we report a novel variant of the task aimed at dissecting the effects of affect manipulations on perceptual and value computations for decision-making under ambiguity in humans. Participants were instructed to judge which way a Gabor patch (250ms presentation) was leaning. If the stimulus leant one way (e.g. left), pressing the REWard key yielded a monetary WIN whilst pressing the SAFE key failed to acquire the WIN. If it leant the other way (e.g. right), pressing the SAFE key avoided a LOSS whilst pressing the REWard key incurred the LOSS. The size (0–100 UK pence) of the offered WIN and threatened LOSS, and the ambiguity of the stimulus (vertical being completely ambiguous) were varied on a trial-by-trial basis, allowing us to investigate how decisions were affected by differing combinations of these factors. Half the subjects performed the task in a ‘Pleasantly’ decorated room and were given a gift (bag of sweets) prior to starting, whilst the other half were in a bare ‘Unpleasant’ room and were not given anything. Although these treatments had little effect on self-reported mood, they did lead to differences in decision-making. All subjects were risk averse under ambiguity, consistent with the notion of loss aversion. Analysis using a Bayesian decision model indicated that Unpleasant Room subjects were (‘pessimistically’) biased towards choosing the SAFE key under ambiguity, but also weighed WINS more heavily than LOSSes compared to Pleasant Room subjects. These apparently contradictory findings may be explained by the influence of affect on different processes underlying decision-making, and the task presented here offers opportunities for further dissecting such processes.     

Facilitation and interference components in the Simon effect.

In four experiments we investigated whether the Simon effect consists of both facilitation and interference. In Experiment 1 subjects had to press one of two lateralized keys in response to one of two stimuli (a rectangle or a square). The stimuli were presented at fixation or to the right or left of it. The stimulus-response mappings could be corresponding (i.e., right key--rectangle or square on the right side), non-corresponding (i.e., right key--rectangle or square on the left side), or neutral (i.e., stimulus in the center). Results showed both facilitation and interference effects, but interference was three times greater than facilitation. To test whether the neutral position was favored by visual acuity, Experiments 2, 3 and 4 used slight different displays where visual acuity was the same at every stimulus position. Results indicated that the Simon effect is comprised of facilitation and interference components of the same magnitude. These findings were discussed with reference to dual-route models of the Simon effect.     

Context-dependent model-based biases in cultural transmission: children's imitation is affected by model age over model knowledge state

Many animals, including humans, acquire information through social learning. Although such information can be acquired easily, its potential unreliability means it should not be used indiscriminately. Cultural ‘transmission biases’ may allow individuals to weigh their reliance on social information according to a model's characteristics. In one of the first studies to juxtapose two model-based biases, we investigated whether the age and knowledge state of a model affected the fidelity of children's copying. Eighty-five 5-year-old children watched a video demonstration of either an adult or child, who had professed either knowledge or ignorance regarding a tool-use task, extracting a reward from that task using both causally relevant and irrelevant actions. Relevant actions were imitated faithfully by children regardless of the model's characteristics, but children who observed an adult reproduced more irrelevant actions than those who observed a child. The professed knowledge state of the model showed a weaker effect on imitation of irrelevant actions. Overall, children favored the use of a ‘copy adults’ bias over a ‘copy task-knowledgeable individual’ bias, even though the latter could potentially have provided more reliable information. The use of such social learning strategies has significant implications for understanding the phenomenon of imitation of irrelevant actions (overimitation), instances of maladaptive information cascades, and cumulative culture.     

Spatial code interference on directional responses.

The interference from an irrelevant position cue was compared in a reaction-time paradigm using voice and manual responses. The subjects were required to say 'left' or 'right' or to press left or right keys in response to arrow directions, and the arrows were presented at left or right side display positions irrelevant to the task. Display position significantly increased latency when it did not match the response to the relevant direction cue for both spatial (key-press) and non-spatial (voice) responses (73 and 59 ms, respectively). When presented alone, the position cue was processed faster than the direction cue for both manual and verbal responses. Results are discussed in terms of a common abstract mediator for left-right responses between modes and the processing speed difference between the relevant and irrelevant cue. The irrelevant left-right position code may occupy some limited-capacity channel ahead of the left-right code derived from processing the relevant direction cue.     

Momentary and global maximizing

Four adult females responded at a computer console, on three constant probability concurrent variable-interval reinforcement schedules. The subjects were instructed to try to obtain as many reinforcers as possible, but were not given any instructions on how to accomplish this task. Three of the four subjects typically allocated responses to the schedule offering the higher reinforcement probability. These results show that some subjects trying to maximize overall reinforcement may respond in accordance with a momentary maximizing strategy.     

Unilateral Auditory Cortex Lesions Impair or Improve Discrimination Learning of Amplitude Modulated Sounds,Depending on Lesion Side

A fundamental principle of brain organization is bilateral symmetry of structures and functions. For spatial sensory and motor information processing, this organization is generally plausible subserving orientation and coordination of a bilaterally symmetric body. However, breaking of the symmetry principle is often seen for functions that depend on convergent information processing and lateralized output control, e.g. left hemispheric dominance for the linguistic speech system. Conversely, a subtle splitting of functions into hemispheres may occur if peripheral information from symmetric sense organs is partly redundant, e.g. auditory pattern recognition, and therefore allows central conceptualizations of complex stimuli from different feature viewpoints, as demonstrated e.g. for hemispheric analysis of frequency modulations in auditory cortex (AC) of mammals including humans. Here we demonstrate that discrimination learning of rapidly but not of slowly amplitude modulated tones is non-uniformly distributed across both hemispheres: While unilateral ablation of left AC in gerbils leads to impairment of normal discrimination learning of rapid amplitude modulations, right side ablations lead to improvement over normal learning. These results point to a rivalry interaction between both ACs in the intact brain where the right side competes with and weakens learning capability maximally attainable by the dominant left side alone.     

Neurons in area V4 of the macaque translate attended visual features into behaviorally relevant categories

Neural processing at most stages of the primate visual system is modulated by selective attention, such that behaviorally relevant information is emphasized at the expenses of irrelevant, potentially distracting information. The form of attention best understood at the cellular level is when stimuli at a given location in the visual field must be selected (space-based attention). In contrast, fewer single-unit recording studies have so far explored the cellular mechanisms of attention operating on individual stimulus features, specifically when one feature (e.g., color) of an object must guide behavioral responses while a second feature (e.g., shape) of the same object is potentially interfering and therefore must be ignored. Here we show that activity of neurons in macaque area V4 can underlie the selection of elemental object features and their "translation" into a categorical format that can directly contribute to the control of the animal's behavior.     

Cognitive components of color vision in honey bees: how conditioning variables modulate color learning and discrimination

Since the demonstration of color vision in honey bees 100 years ago by Karl von Frisch, appetitive conditioning to color targets has been used as the principal way to access behavioral aspects of bee color vision. Yet, analyses on how conditioning parameters affect color perception remained scarce. Conclusions on bee color vision have often been made without referring them to the experimental context in which they were obtained, and thus presented as absolute facts instead of realizing that subtle variations in conditioning procedures might yield different results. Here, we review evidence showing that color learning and discrimination in bees are not governed by immutable properties of their visual system, but depend on how the insects are trained and thus learn a task. The use of absolute or differential conditioning protocols, the presence of aversive reinforcement in differential conditioning and the degrees of freedom of motor components determine dramatic variations in color discrimination. We, thus, suggest top-down attentional modulation of color vision to explain the changes in color learning and discrimination reviewed here. We discuss the possible neural mechanisms of this modulation and conclude that color vision experiments require a careful consideration of how training parameters shape behavioral responses.     

The additive factor method in rat information processing

Sternberg introduced the additive factor method as a tool for discovering and studying in choice reaction time situations in human subjects. Underlying the method is the notion that if information processing consists of successive stages, then different factors affecting different stages must have additive effects on reaction-time measurements. The additive factor method has been extensively used in humans but few attempts have been made to examine whether or not additive reaction time patterns were obtainable in rats. In the present work, the subjects (13 Long–Evans rats) had to press down a lever and to respond to a visual (right or left) stimulus by releasing the lever and introducing the head into a lateral (right or left) operandum. The present findings show an additive pattern of effects of signal luminance and foreperiod duration on the mean reaction time which suggests that we have successfully manipulated two stages of rat information processing — the preprocessing stage and the motor adjustment stage, respectively. Accordingly, the present study sets the basis for the basis for investigating the effects of invasive manipulations (pharmacological and/or lesional) on information processing stages.     

Effects of sleep loss and circadian rhythm on executive inhibitory control in the Stroop and Simon tasks

This study assessed the influence of sleep loss and circadian rhythm on executive inhibitory control (i.e., the ability to inhibit conflicting response tendencies due to irrelevant information). Twelve ordinarily diurnally active, healthy young male participants performed the Stroop and the Simon task every 3?h in a 40-h constant routine protocol that comprised constant wakefulness under controlled behavioral and environmental conditions. In both tasks, overall performance showed clear circadian rhythm and sleep-loss effects. However, both Stroop and Simon interference remained unchanged across the 40?h of wakefulness, suggesting that neither cumulative sleep loss nor the circadian clock affects executive inhibitory control. The present findings challenge the widely held view that executive functions are especially vulnerable to the influence of sleep loss and circadian rhythm.     

Right medial temporal-lobe contribution to object-location memory.

An important aspect of normal human memory, and one humans share with many other species, is the ability to remember the location of objects in their environment. There is by now strong evidence from the study of epileptic patients undergoing brain surgery that right temporal-lobe lesions that encroach extensively upon the hippocampal and parahippocampal gyrus impair the delayed, but not the immediate, recall of the location of objects within a random array. These findings have now been extended to a multiple-trial, spatial-array learning task; by including not only patients tested after unilateral anterior temporal lobectomy but also those with a selective left or right amygdalohippocampectomy, it has been shown that the deficits associated with right hippocampal lesions are not dependent upon conjoint damage to the lateral temporal neocortex. Furthermore, the fact that on the learning task no group differences were seen on Trial 1, at zero delay, strengthened the view that the impairment was in the maintenance and subsequent retrieval of information rather than in its initial encoding. These results left unresolved the question of whether the deficit was in the mediation of object-place associations or whether it could be reduced to a more general impairment in memory for location as such. Also left unanswered was the neuroanatomical question as to the relative contributions of the hippocampus and the parahippocampal gyrus to the performance of the experimental tasks. These questions were addressed in two blood-flow activation studies that made use of positron emission tomography (PET) and magnetic resonance imaging (MRI) and incorporated computerized versions of object-location and simple-location memory tasks. Taken together, the results point to a special contribution from the anterior part of the right parahippocampal gyrus, probably corresponding to the entorhinal cortex, to the retrieval of object-place associations, a result consonant with neurophysiological findings in non-human primates.     

Virtual dyscalculia induced by parietal-lobe TMS impairs automatic magnitude processing

People suffering from developmental dyscalculia encounter difficulties in automatically accessing numerical magnitudes [1-3]. For example, when instructed to attend to the physical size of a number while ignoring its numerical value, dyscalculic subjects, unlike healthy participants, fail to process the irrelevant dimension automatically and subsequently show a smaller size-congruity effect (difference in reaction time between incongruent [e.g., a physically large 2 and a physically small 4] and congruent [e.g., a physically small 2 and a physically large 4] conditions), and no facilitation (neutral [e.g., a physically small 2 and a physically large 2] versus congruent) [3]. Previous imaging studies determined the intraparietal sulcus (IPS) as a central area for numerical processing [4-11]. A few studies tried to identify the brain dysfunction underlying developmental dyscalculia but yielded mixed results regarding the involvement of the left [12] or the right [13] IPS. Here we applied fMRI-guided TMS neuronavigation to disrupt left- or right-IPS activation clusters in order to induce dyscalculic-like behavioral deficits in healthy volunteers. Automatic magnitude processing was impaired only during disruption of right-IPS activity. When using the identical paradigm with dyscalculic participants, we reproduced a result pattern similar to that obtained with nondyscalculic volunteers during right-IPS disruption. These findings provide direct evidence for the functional role of right IPS in automatic magnitude processing.