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A viewpoint-independent process for spatial reorientation.
Reorientation tasks, in which disoriented participants attempt to relocate objects using different visual cues, have previously been understood to depend on representing aspects of the global organisation of the space, for example its major axis for judgements based on geometry. Careful analysis of the visual information available for these tasks shows that successful performance could be based on the much simpler process of storing a visual 'snapshot' at the target location, and subsequently moving in order to match it. We tested 4-8-year olds on a new spatial reorientation task that could not be solved based on information directly contained in any retinal projection that they had been exposed to, but required participants to infer how the space is structured. Only 6-8-year olds showed flexible recall from novel viewpoints. Five-year olds were able to recall locations given movement information or a unique proximal landmark, but without these they could not do so, even when they were not disoriented or when the landmark was a familiar object. These results indicate that early developing spatial abilities based on view matching and self motion are supplemented by a later-developing process that takes into account the structure of spatial layouts and so enables flexible recall from arbitrary viewpoints.
Orientation-reversal and phase-reversal visual evoked potentials in full-term infants with brain lesions: a longitudinal study.
The onset and maturation of visual cortical mechanisms can be recorded by using steady-state visual evoked potentials. The aim of this study was to evaluate and compare orientation-reversal (OR) and phase-reversal (PH) VEP as indicators of the maturation of cortical function in a population of fullterm infants with brain lesions on neonatal MRI. Forty-six infants with brain lesions on neonatal MRI were tested on both PH and OR VEP at 8 reversals/second at the age of 5 months and, if the responses were not significant, at a lower temporal frequency (4 reversals/second). Children whose VEPs were not significant at 5 months were tested longitudinally at 6, 9, 12 and 18 months. The results showed that 23 of the 46 infants (50%) did not show significant responses at 5 months and that while in 7 of the 23 (14% of the whole cohort) the responses became significant between 5 and 12 months, in the other 16 infants (34%) the VEP responses were persistently abnormal. Children with focal lesions, such as focal infarction or haemorrhages, tended to show normal or only mildly delayed VEP while more generalised lesions, such as the ones seen in infants with hypoxic-ischaemic encephalopathy grade 2 and 3, tended to be associated with abnormal VEP responses. The involvement of the optic radiations and occipital cortex was not always associated with abnormal VEP responses but the concomitant involvement of the basal ganglia was always associated with abnormal VEP. We were also able to demonstrate that VEP can be also used as a prognostic indicator: while normal OR VEP are reliably associated with a normal visual and neurodevelopmental outcome, abnormal 4 OR or 8 PH at 5 months are consistently associated with abnormal outcome.
Integration across directions in dynamic random dot displays: vector summation or winner take all?
Recent studies have clearly demonstrated that the activity of directionally selective neuronal populations in the middle temporal (MT) and medial superior temporal (MST) cortical areas plays a direct role in the judgment of the direction of visual motion. However, the way in which the information is derived from a population of neurons remains unknown. Two principal models have been suggested in the past: the vector summation model suggests that the responses of neurons encoding all directions of motion are weighted and pooled to obtained an accurate estimate of the mean direction of motion; the winner-take-all model is based on a competition between different direction-specific channels, so that decisions are cast in favor of the channel generating the strongest directional signal. To discriminate between these two models we generated random dot stimuli that contained an asymmetric distribution of directions of motion. Human subjects were asked to adjust the global direction of motion to the upward vertical direction. When the directional signals were of similar strength, subjects tended to perceive global motion in the mean direction of motion (corresponding to vector summation), but as one directional signal became more prominent, most subjects' settings diverged from the mean towards the modal direction of motion. Some subjects could either match the mean or the modal direction of motion in the display, depending on the task instructions. These results suggest that the perceptual judgment of direction of motion is not based on any rigid algorithm generating a single valued output. Rather, human observers are able to judge different aspects of the distribution of activity in a cortical area depending on the task requirements.
What is noise for the motion system?
Motion coherence thresholds in random-dot patterns have been widely adopted as a measure of performance in visual motion processing. However, there has been diversity in the type of "noise" in which a coherent motion signal has to be detected. Here we compare coherence thresholds for three ways of creating motion noise: dots replotted in random positions in each new frame; dots with a set displacement but following a random walk from frame to frame; or dots moving in random directions which remain constant for a given dot over a sequence of displacements. In each case, the signal dots may either remain the same throughout the display sequence, or the signal dots may be re-selected afresh on each frame ("different"). With our display (3 deg square, 120 msec exposure, velocity = 5 or 10 deg sec-1), all these different noise conditions yielded similar thresholds around 5-8%. There were some small but systematic differences between conditions. Thresholds in random-direction displays were consistently higher than those in random-walk or random-position displays, especially at the lower velocity. However, this effect is much smaller than would be expected from the increased standard error of the noise mean in random direction, perhaps because the motion system integrates information most effectively over a local region of space and/or time. Subjects" performance could not be explained by a strategy of identifying individual signal dots with extended trajectories. The similarity between random-walk and random-position thresholds implies that subjects do not exploit the marked differences in speed distribution between signal and noise dots in the latter case. The practical message for the design and interpretation of experiments using coherence thresholds is that the results are not much affected by the choice of noise, at least within the range of stimuli tested here.
Contrast sensitivity function of preschool children.
A procedure specifically adapted for children of preschool age has been used to measure contrast sensitivity in emmetropic children aged 3 to 5 years. Mothers of the children acted as adult observers using the same procedure. The results show that the contrast sensitivity function of adults and children is very similar, sensitivity for the children being slightly lower than that for adults at all spatial frequencies. The sensory and cognitive factors involved in these differences are discussed.
Does size matter? Subsegmental cues to vowel mispronunciation detection.
Children look longer at a familiar object when presented with either correct pronunciations or small mispronunciations of consonants in the object's label, but not following larger mispronunciations. The current article examines whether children display a similar graded sensitivity to different degrees of mispronunciations of the vowels in familiar words, by testing children's sensitivity to 1-feature, 2-feature and 3-feature mispronunciations of the vowels of familiar labels: Children aged 1 ; 6 did not show a graded sensitivity to vowel mispronunciations, even when the trial length was increased to allow them more time to form a response. Two-year-olds displayed a robust sensitivity to increases in vowel mispronunciation size, differentiating between small and large mispronunciations. While this suggests that early lexical representations contain information about the features contributing to vocalic identity, we present evidence that this graded sensitivity is better explained by the acoustic characteristics of the different mispronunciation types presented to children.
A large-scale distributed network for covert spatial attention: further anatomical delineation based on stringent behavioural and cognitive controls.
Functional MRI was used to examine cerebral activations in 12 subjects while they performed a spatial attention task. This study applied more stringent behavioural and cognitive controls than previously used for similar experiments: (i) subjects were included only if they showed evidence of attentional shifts while performing the task in the magnet; (ii) the experimental task and baseline condition were designed to eliminate the contributions of motor output, visual fixation, inhibition of eye movements, working memory and the conditional (no-go) component of responding. Activations were seen in all three hypothesized cortical epicentres forming a network for spatial attention: the lateral premotor cortex (frontal eye fields), the posterior parietal cortex and the cingulate cortex. Subcortical activations were seen in the basal ganglia and the thalamus. Although the task required attention to be equally shifted to the left and to the right, eight of 10 subjects showed a greater area of activation in the right parietal cortex, consistent with the specialization of the right hemisphere for spatial attention. Other areas of significant activation included the posterior temporo-occipital cortex and the anterior insula. The temporo-occipital activation was within a region broadly defined as MT+ (where MT is the middle temporal area) which contains the human equivalent of area MT in the macaque monkey. This temporo-occipital area appears to constitute a major component of the functional network activated by this spatial attention task. Its activation may reflect the 'inferred' shift of the attentional focus across the visual scene.
Acquisition of the temporal and ordinal structure of movement sequences in incidental learning.
We investigated the acquisition and integration of temporal and ordinal sequence information in an incidental learning model of motor skill acquisition (the serial reaction time task). Human participants were exposed to a stimulus-response sequence that had temporal structure, ordinal structure, or both. By changing the temporal or ordinal structure, or both, we were able to ask two questions: first, does a regular temporal structure facilitate learning of an ordinal sequence and second, is a temporal sequence, presented in the context of a random ordinal sequence of finger movements, "picked up" through incidental learning? We found that a predictable temporal structure greatly facilitated the learning of an ordinal sequence but was not learned when presented in isolation. The results suggest that when motor skills are acquired under incidental learning conditions, timing is represented at a level specific to the ordinal sequence of movements rather than as an independent temporal template.
Hunger selectively modulates corticolimbic activation to food stimuli in humans.
Functional magnetic resonance imaging (fMRI) was used to determine whether visual responses to food in the human amygdala and related corticolimbic structures would be selectively altered by changes in states of hunger. Participants viewed images of motivationally relevant (food) and motivationally irrelevant (tool) objects while undergoing fMRI in alternately hungry and satiated conditions. Food-related visual stimuli elicited greater responses in the amygdala, parahippocampal gyrus. and anterior fusiform gyrus when participants were in a hungry state relative to a satiated state. The state-dependent activation of these brain structures did not generalize to the motivationally irrelevant objects. These results support the hypothesis that the amygdala and associated inferotemporal regions are involved in the integration of subjective interoceptive states with relevant sensory cues processed along the ventral visual stream.
The large-scale neural network for spatial attention displays multifunctional overlap but differential asymmetry.
Functional magnetic resonance imaging (fMRI) was used to determine the brain regions activated by two types of covert visuospatial attentional shifts: one based on exogenous spatial priming and the other on foveally presented cues which endogenously regulated the direction of spatial expectancy. Activations were seen in the cortical and subcortical components of a previously characterized attentional network, namely, the frontal eye fields, posterior parietal cortex, the cingulate gyrus, the putamen, and the thalamus. Additional activations occurred in the anterior insula, dorsolateral prefrontal cortex, temporo-occipital cortex in the middle and inferior temporal gyri, the supplementary motor area, and the cerebellum. Direct comparisons showed a nearly complete overlap in the location of activations resulting from the two tasks. However, the spatial priming task displayed a more pronounced rightward asymmetry of parietal activation, and a conjunction analysis showed that the area of posterior parietal cortex jointly activated by both tasks was more extensive in the right hemisphere. Furthermore, the posterior parietal and temporo-occipital activations were more pronounced in the task of endogenous attentional shifts. The results show that both exogenous (based on spatial priming) and endogenous (based on expectancy cueing) shifts of attention are subserved by a common network of cortical and subcortical regions. However, the differences between the two tasks, especially in the degree of rightward asymmetry, suggests that the pattern of activation within this network may show variations that reflect the specific attributes of the attentional task.
Reward associations magnify memory-based biases on perception.
Long-term spatial contextual memories are a rich source of predictions about the likely locations of relevant objects in the environment and should enable tuning of neural processing of unfolding events to optimize perception and action. Of particular importance is whether and how the reward outcome of past events can impact perception. We combined behavioral measures with recordings of brain activity with high temporal resolution to test whether the previous reward outcome associated with a memory could modulate the impact of memory-based biases on perception, and if so, the level(s) at which visual neural processing is biased by reward-associated memory-guided attention. Data showed that past rewards potentiate the effects of spatial memories upon the discrimination of target objects embedded within complex scenes starting from early perceptual stages. We show that a single reward outcome of learning impacts on how we perceive events in our complex environments.
Age-related changes in orienting attention in time.
Temporal cues guide attentional resources toward relevant points in time, resulting in optimized behavioral performance. Although deficits in aspects of attention have been documented in older adults, it remains unknown whether the critical ability to orient attention in time is affected by normal aging. To address this, younger and older adults participated in a temporally cued target-response experiment while electroencephalographic data were recorded. Three conditions (one detection and two discrimination tasks) were used to manipulate task complexity. Response times show that younger adults, but not older adults, used temporal cues to enhance performance regardless of task complexity. Similarly, alpha band activity (8-12 Hz) and the contingent negative variation preceding targets indicated that only younger adults engaged prestimulus, anticipatory neural mechanisms associated with temporal cues. Overall, these results provide novel evidence that older adults do not use temporal cues to orient attention in time and support an expectation deficit in normal aging.
Modelling distractor devaluation (DD) and its neurophysiological correlates.
A series of recent studies have shown that selective attention can influence the emotional value of both selected as well as ignored items. Specifically, ignored items (distractors) were consistently rated less positively in emotional evaluations, following attentional selection, relative to (typically) simultaneously presented items (targets). Furthermore, a known electrophysiological index of attentional selectivity (N2pc) was shown to correlate with the magnitude of the observed 'distractor devaluation' (DD). A neural model is presented here to account for these findings by means of a plausible mechanism linking attentional processes to emotional evaluations. This mechanism relies on the transformation of attentional inhibition of the distractor into a reduction of the value of that distractor. The model is successful in reproducing the existent behavioural results as well as the observed link between the magnitude of the attentional N2pc and the magnitude of DD. Moreover, the model proposes a series of testable hypotheses as well as specific predictions that call for further experimental investigation.
Temporal expectation enhances contrast sensitivity by phase entrainment of low-frequency oscillations in visual cortex.
Although it is increasingly accepted that temporal expectation can modulate early perceptual processing, the underlying neural computations remain unknown. In the present study, we combined a psychophysical paradigm with electrophysiological recordings to investigate the putative contribution of low-frequency oscillatory activity in mediating the modulation of visual perception by temporal expectation. Human participants judged the orientation of brief targets (visual Gabor patterns tilted clockwise or counterclockwise) embedded within temporally regular or irregular streams of noise-patches used as temporal cues. Psychophysical results indicated that temporal expectation enhanced the contrast sensitivity of visual targets. A diffusion model indicated that rhythmic temporal expectation modulated the signal-to-noise gain of visual processing. The concurrent electrophysiological data revealed that the phase of delta oscillations overlying human visual cortex (1-4 Hz) was predictive of the quality of target processing only in regular streams of events. Moreover, in the regular condition, the optimum phase of these perception-predictive oscillations occurred in anticipation of the expected events. Together, these results show a strong correspondence between psychophysical and neurophysiological data, suggesting that the phase entrainment of low-frequency oscillations to external sensory cues can serve as an important and flexible mechanism for enhancing sensory processing.
The cerebellum predicts the timing of perceptual events.
Prospective (forward) temporal-spatial models are essential for both action and perception, but the literature on perceptual prediction has primarily been limited to the spatial domain. In this study we asked how the neural systems of perceptual prediction change, when change-over-time must be modeled. We used a naturalistic paradigm in which observers had to extrapolate the trajectory of an occluded moving object to make perceptual judgments based on the spatial (direction) or temporal-spatial (velocity) characteristics of object motion. Using functional magnetic resonance imaging we found that a region in posterior cerebellum (lobule VII crus 1) was engaged specifically when a temporal-spatial model was required (velocity judgment task), suggesting that circuitry involved in motor forward-modeling may also be engaged in perceptual prediction when a model of change-over-time is required. This cerebellar region appears to supply a temporal signal to cortical networks involved in spatial orienting: a frontal-parietal network associated with attentional orienting was engaged in both (spatial and temporal-spatial) tasks, but functional connectivity between these regions and the posterior cerebellum was enhanced in the temporal-spatial prediction task. In addition to the oculomotor spatial orienting network, regions involved in hand movements (aIP and PMv) were recruited in the temporal-spatial task, suggesting that the nature of perceptual prediction may bias the recruitment of sensory-motor networks in orienting. Finally, in temporal-spatial prediction, functional connectivity was enhanced between the cerebellum and the putamen, a structure which has been proposed to supply the brain's metric of time, in the temporal-spatial prediction task.
Temporal expectation improves the quality of sensory information.
It is increasingly clear that we extract patterns of temporal regularity between events to optimize information processing. Whereas some of the mechanisms for facilitating action preparation and execution have been well documented, much less is understood about whether and how temporal expectations influence visual perception. We used a psychophysical paradigm and computational modeling to investigate the mechanisms by which temporal expectation can modulate visual perception. Visual targets appeared in a stream of noise-patches separated by a fixed (400 ms regular condition) or jittered (200/300/400/500/600 ms irregular condition) intervals. Targets were visual gratings tilted 45° clockwise or counter-clockwise, presented at one of seven contrast levels. Human observers were required to perform an orientation discrimination (i.e., left or right). Psychometric functions for contrast sensitivity fitted for the regular and irregular conditions indicated that temporal expectation modulates perceptual processing by enhancing the contrast sensitivity of visual targets. This increase in the signal strength was accompanied by a reduction in reaction times. A diffusion model indicated that rhythmic temporal expectation enhanced the signal-to-noise gain of the sensory evidence upon which decisions were made. These effects support the idea that temporal structure of external events can entrain the attentional focus and psychophysical data, optimizing the processing of relevant sensory information.
Synergistic effect of combined temporal and spatial expectations on visual attention.
We developed a naturalistic behavioral task to investigate the influence of spatial and temporal expectations on attentional orienting to moving targets. In this task, the movement of an object before its disappearance under an occluding barrier generated expectations concerning the location and/or time of its reappearance. Four different trial types were intermixed, each inducing a different state of expectation: no expectation, only spatial expectation about the location of reappearance, only temporal expectation about the moment of reappearance, and combined spatial and temporal expectation. The behavioral validity of the task was shown by the fact that all expectation conditions produced significantly shorter reaction times than the control state of no expectation. Spatial attention modulated early perceptual analysis in extrastriate areas, as demonstrated by significant enhancement of the visual P1 component. Temporal attention alone had no effect on P1 but instead modulated response-specific components. However, when spatial and temporal attention were combined, the enhancement of perceptual processing was significantly augmented, leading to a greater enhancement of the P1 component than by spatial attention alone. Perceptual analysis reflected by the P1 component correlated significantly with reaction times. In summary, event-related potentials revealed the presence of individual modulatory effects attributable to spatial and temporal expectation as well as synergistic effects indicative of an interaction of the two. This synergistic effect is likely to play a critical role in directing attention to the reappearance of a temporarily occluded moving target, a process of obvious importance in everyday situations.
α oscillations related to anticipatory attention follow temporal expectations.
Temporal expectations have been shown to enhance visual analysis of task-relevant events, especially when these are coupled with spatial expectations. Oscillatory brain activity, particularly in the alpha band, has been implicated in regulating excitability in visual areas as a function of anticipatory spatial attention. Here we asked whether temporal expectations derived from regular, rhythmic events can modulate ongoing oscillatory alpha-band activity, so that the changes in cortical excitability are focused over the time intervals at which target events are expected. The task we used involved making a perceptual discrimination about a small target stimulus that reappeared from "behind" a peripheral occluding band. Temporal expectations were manipulated by the regular, rhythmic versus irregular, arrhythmic approach of the stimulus toward the occluding band. Alpha-band activity was measured during the occlusion period, in which no stimulus was presented, but target reappearance was anticipated in conditions of high versus low temporal expectation. Time-frequency analysis showed that the amplitude of alpha-desynchronization followed the time course of temporal expectations. Alpha desynchronization increased rhythmically, peaking just before the expected reappearance of target times. Analysis of the event-related potentials evoked by the subsequent target stimuli showed enhancement of processing at both visual and motor stages. Our findings support a role for oscillations in regulating cortical excitability and suggest a plausible mechanism for biasing perception and action by temporal expectations.