Priming of motion direction and area V5/MT: a test of perceptual memory

Presentation of supraliminal or subliminal visual stimuli that can (or cannot) be detected or identified can improve the probability of the same stimulus being detected over a subsequent period of seconds, hours or longer. The locus and nature of this perceptual priming effect was examined, using suprathreshold stimuli, in subjects who received repetitive pulse transcranial magnetic stimulation over the posterior occipital cortex, the extrastriate motion area V5/MT or the right posterior parietal cortex during the intertrial interval of a visual motion direction discrimination task. Perceptual priming observed in a control condition was abolished when area V5/MT was stimulated but was not affected by magnetic stimulation over striate or parietal sites. The effect of transcranial magnetic stimulation (TMS) on priming was specific to site (V5/MT) and to task - colour priming was unaffected by TMS over V5/MT. The results parallel, in the motion domain, recent demonstrations of the importance of macaque areas V4 and TEO for priming in the colour and form domains.     

Flexible, capacity-limited activity of posterior parietal cortex in perceptual as well as visual short-term memory tasks

It has recently been shown, using functional magnetic resonance imaging with a change detection paradigm, that activity in posterior parietal cortex (PPC) correlates with the limited number of objects held in visual short-term memory (VSTM). We replicate this finding and extend it to tasks that use similar stimuli, but without explicit memory requirements. As well as a perceptual task used previously (detecting an item at fixation), 2 additional tasks were designed to increase attentional demands across space (searching for a red item anywhere in the array) and across both space and time (detecting a staggered offset after prolonged viewing of the array). During the VSTM task, a capacity-limited set-size effect was seen in PPC as well as occipital and frontal regions. However, the PPC showed similar activity during 2 of the tasks not requiring VSTM. These findings cannot easily be explained by behavioral performance measures or memory demands alone, suggesting a role of the PPC in processing a limited number of discrete object representations, whether in the current perceptual scene or working memory. The differential influence of item load across perceptual tasks is consistent with task requirements affecting the form of these representations.     

The perceptual and functional consequences of parietal top-down modulation on the visual cortex

The posterior parietal cortex (PPC) has been proposed to play a critical role in exerting top-down influences on occipital visual areas. By inducing activity in the PPC (angular gyrus) using transcranial magnetic stimulation (TMS), and using the phosphene threshold as a measure of visual cortical excitability, we investigated the functional role of this region in modulating the activity of the visual cortex. When triple-pulses of TMS were applied over the PPC unilaterally, the intensity of stimulation required to elicit a phosphene from the visual cortex (area V1/V2) was reduced, indicating an increase in visual cortical excitability. The increased excitability that was observed with unilateral TMS was abolished when TMS was applied over the PPC bilaterally. Our results provide a demonstration of the top-down modulation exerted by the PPC on the visual cortex and show that these effects are subject to interhemispheric competition.     

Working memory specific activity in auditory cortex: potential correlates of sequential processing and maintenance

Working memory (WM) tasks involve several interrelated processes during which past information must be transiently maintained, recalled, and compared with test items according to previously instructed rules. It is not clear whether the rule-specific comparisons of perceptual with memorized items are only performed in previously identified frontal and parietal WM areas or whether these areas orchestrate such comparisons by feedback to sensory cortex. We tested the latter hypothesis by focusing on auditory cortex (AC) areas with low-noise functional magnetic resonance imaging in a 2-back WM task involving frequency-modulated (FM) tones. The control condition was a 0-back task on the same stimuli. Analysis of the group data identified an area on right planum temporale equally activated by both tasks and an area on the left planum temporale specifically involved in the 2-back task. A region of interest analysis in each individual revealed that activation on the left planum temporale in the 2-back task positively correlated with the task performance of the subjects. This strongly suggests a prominent role of the AC in 2-back WM tasks. In conjunction with previous findings on FM processing, the left lateralized effect presumably reflects the complex sequential processing demand of the 2-back matching to sample task.     

The planning and guiding of reading saccades: a repetitive transcranial magnetic stimulation study.

A previous positron emission tomography study that investigated the cortical areas involved in directing eye movements during text reading showed two areas of extra-occipital asymmetry: left > right posterior parietal cortex (PPC), and right > left frontal eye-field (FEF). We used the temporal resolution of repetitive TMS (rTMS) to isolate the contributions of the left and right PPC and FEF to the planning and execution of rightward reading saccades. We present eye-movement data collected during text reading, which involves the initiation and maintenance of a series of saccades (scanpath). rTMS over the left but not right PPC slowed reading speeds for the whole array of words, indicating that this area is involved throughout the scanpath. rTMS over the right but not the left FEF slowed the time to make the first saccade, but only when triggered before the stimuli appeared, demonstrating that the role of this region is in the preparation of the scanpath. Our results are compatible with the hypotheses that the left PPC maintains reading saccades along a line of text while the right FEF is involved in the preparation of the motor plan for the scanpath at the start of each new line of text.     

Double dissociation of V1 and V5/MT activity in visual awareness

The critical time windows of the contribution of V1 and V5/MT to visual awareness of moving visual stimuli were compared by administering transcranial magnetic stimulation (TMS) to V1 or V5/MT in various time intervals from stimulus offset during performance of a simple motion detection task. Our results show a double dissociation in which the critical period of V1 both predates and postdates that of V5/MT, and where stimulation of either V1 at V5/MT's critical period or V5/MT at V1's critical period does not impair performance. These findings demonstrate the importance of back-projections from V5/MT to V1 in awareness of real motion stimuli.     

The multiple roles of visual cortical areas MT/MST in remembering the direction of visual motion

Although the role of cortical areas MT and MST (MT/MST) in the processing of directional motion information is well established, little is known about the way these areas contribute to the execution of complex behavioral tasks requiring the use of such information. We tested monkeys with unilateral lesions of these areas on a visual working memory task in which motion signals not only had to be encoded, but also stored for brief periods of time and then retrieved. The monkeys compared the directions of motion of two random-dot stimuli, sample and test, separated by a temporal delay. By increasing the temporal delay and spatially separating the two stimuli, placing one in the affected visual field and the other in the intact visual field, we were able to assess the contribution of MT/MST to specific components of the task: encoding (sample), retention (delay) and encoding/retrieval/comparison (test). We found that the effects of MT/MST lesions on specific components depended upon the demands of the task and the nature of the visual motion stimuli. Whenever stimuli consisted of random dots moving in a broad range of directions, MT/MST lesions appeared to affect encoding. Furthermore, when the lesions affected encoding of the sample, retention of the direction of stimulus motion was also affected. However, when the stimulus was coherent and the emphasis of the task was on the comparison of small direction differences, the absence of MT/MST had major impact on the retrieval/comparison component of the task and not on encoding or storage.     

Visual area V5/MT remembers "what" but not "where"

Priming for motion direction has been shown to depend upon the functional integrity of extrastriate area V5/MT. Its retinotopic organization and the interactions recently found between motion adaptation and misperceived localization may suggest, for this area, a role for priming of spatial position in addition to the established priming of motion direction. Disruption of V5/MT with repetitive transcranial magnetic stimulation during the intertrial interval had the effect of abolishing priming of motion direction but no effect in priming of spatial position. These effects cannot be explained in terms of perception or task demands but only in terms of the effects of information irrelevant to the correct performance of the task stored over the intertrial interval. We suggest that the attribute of spatial position might be stored in short-term memory either in earlier areas of the motion pathways such as V3 or in higher cortical areas traditionally associated with the analysis of spatial information, for example, posterior parietal cortex or the frontal eye fields.     

The contribution of the human FEF and SEF to smooth pursuit initiation

Smooth pursuit eye movements function to keep moving targets foveated. Behavioral studies have shown that pursuit is particularly effective for predictable target motion. There is evidence that both the frontal eye field (FEF) and supplementary eye field (SEF) (also known as the dorsomedial frontal cortex) contribute to pursuit control. The goal of the current experiment was to determine whether these 2 areas made different contributions to the initiation of pursuit in response to predictable compared with unpredictable target motion. Transcranial magnetic stimulation (TMS) was used in 5 healthy human participants to temporarily disrupt each area around the time of target motion onset. TMS over the FEF delayed contraversive pursuit markedly more than ipsiversive pursuit and this direction-dependent difference was more deeply modulated during pursuit of unpredictable than predictable target motion. By contrast, TMS over the SEF resulted in a much more muted modulation of pursuit latency that was similar across both predictable and unpredictable conditions. Taken together, we conclude that the human FEF, but not the SEF, makes a significant contribution to the processing required during the preparation of contraversive pursuit responses to unpredictable target motion and this contribution is less vital during pursuit to predictable target motion.     

Activation of area MT/V5 and the right inferior parietal cortex during the discrimination of transient direction changes in translational motion

The perception of changes in the direction of objects that translate in space is an important function of our visual system. Here we investigate the brain electrical phenomena underlying such a function by using a combination of magnetoencephalography (MEG) and magnetic resonance imaging. We recorded MEG-evoked responses in 9 healthy human subjects while they discriminated the direction of a transient change in a translationally moving random dot pattern presented either to the right or to the left of a central fixation point. We found that responses reached their maximum in 2 main regions corresponding to motion processing area middle temporal (MT)/V5 contralateral to the stimulated visual field, and to the right inferior parietal lobe (rIPL). The activation latencies were very similar in both regions ( approximately 135 ms) following the direction change onset. Our findings suggest that area MT/V5 provides the strongest sensory signal in response to changes in the direction of translational motion, whereas area rIPL may be involved either in the sensory processing of transient motion signals or in the processing of signals related to orienting of attention.     

Role of the prefrontal cortex in the foreperiod effect: TMS evidence for dual mechanisms in temporal preparation

The involvement of right dorsolateral prefrontal cortex (rDLPFC) in explicit temporal processing is well documented. Conversely, the role of this area in implicit temporal processing (e.g., foreperiod [FP] effect) is still poorly understood. The FP effect, usually observed when a range of variable FPs occur randomly and equiprobably, consists of reaction times (RTs) decreasing as the FP increases. Moreover, in such paradigms, RTs increase as a function of the preceding FP (i.e., sequential effects). Patients with lesions of the rDLPFC do not show the typical FP effect. The present study aimed to replicate these results in healthy adults using transcranial magnetic stimulation (TMS) and to further investigate whether any change of sequential effects follows a reduction of the FP effect. The results of 2 experiments (with simple and choice RT tasks, respectively) indicate that the FP effect was significantly reduced after TMS over the rDLPFC, whereas no effect was observed after stimulation of a left contralateral site and the right angular gyrus. Conversely, sequential effects were not influenced by TMS. A dual-process model of the FP phenomena is proposed to interpret the dissociation found between the 2 effects.     

The kinetic occipital region in human visual cortex

In the present study we showed that the kinetic occipital (KO) region, located laterally in occipital cortex approximately 20 mm behind human MT/V5, can be strongly and bilaterally activated under passive viewing conditions. We used continuous, randomly changing visual stimulation to compare kinetic gratings to uniform motion and kinetic gratings to luminance defined gratings. The KO activations under these passive conditions are stronger than those observed when the two types of gratings are compared under active conditions, i.e. while subjects perform a task (counting gratings of a given orientation). Region KO was shown to process both shape and motion information, the conjunction of which is typically present in kinetic contours. Area MT/V5 also processes these two aspects of visual stimulation but favors motion signals. Clear segregation of shape and motion processing was observed only in occipitotemporal and parietal regions respectively. Although neurons with properties similar to those derived from the conditions activating the KO region have been documented in the macaque monkey, their location seems inappropriate for them to correspond to the KO activation observed in humans.      

Translaminar differentiation of visually guided behaviors revealed by restricted cerebral cooling deactivation

The purpose of the present study was to test the hypothesis that superficial and deep layers within a single cerebral region influence cerebral functions and behaviors in different ways. For this test, we selected posterior middle suprasylvian (pMS) sulcal cortex of the cat, a suspected homolog of the area V5 complex of primates, because the region has been implicated in several visually guided behaviors. Cats were trained on three tasks: (1) discrimination of direction of motion; (2) discrimination of static patterns partially obscured by static or moving masks; and (3) visual detection and orienting. Cooling of cryoloops in contact with pMS sulcal cortex to 8+/-1 degrees C selectively and completely impaired performance on the two motion discrimination tasks (1 and 2), while leaving the detection and orienting task (task 3) unimpaired. Further cooling to 3 degrees C resulted in an additional complete impairment of task 3. The 8 degrees C temperature resulted in silencing of neuronal activity in the supragranular layers (I-III) and the 3 degrees C temperature silenced activity throughout the thickness of pMS sulcal cortex. The variation in behavioral performance with covariation of cryoloop temperature and vertical, but not lateral, spread of deactivation shows that deactivation of superficial cerebral layers alone was sufficient to completely impair performance on the two motion discrimination tasks, whereas additional deactivation of the deep layers was essential to block performance on the detection and orienting task. Thus, these results show a functional bipartite division of labor between upper and lower cortical layers that is supported by efferent connectional anatomy. Similar bipartite division into upper and lower layers may be a general feature of cerebral cortical architecture, signal processing and guidance of behavior.     

Contributions of the PPC to online control of visually guided reaching movements assessed with fMRI-guided TMS

The posterior parietal cortex (PPC) plays an important role in controlling voluntary movements by continuously integrating sensory information about body state and the environment. We tested which subregions of the PPC contribute to the processing of target- and body-related visual information while reaching for an object, using a reaching paradigm with 2 types of visual perturbation: displacement of the visual target and displacement of the visual feedback about the hand position. Initially, functional magnetic resonance imaging (fMRI) was used to localize putative target areas involved in online corrections of movements in response to perturbations. The causal contribution of these areas to online correction was tested in subsequent neuronavigated transcranial magnetic stimulation (TMS) experiments. Robust TMS effects occurred at distinct anatomical sites along the anterior intraparietal sulcus (aIPS) and the anterior part of the supramarginal gyrus for both perturbations. TMS over neighboring sites did not affect online control. Our results support the hypothesis that the aIPS is more generally involved in visually guided control of movements, independent of body effectors and nature of the visual information. Furthermore, they suggest that the human network of PPC subregions controlling goal-directed visuomotor processes extends more inferiorly than previously thought. Our results also point toward a good spatial specificity of the TMS effects.     

Top-down controlled visual dimension weighting: an event-related fMRI study

Target detection in visual singleton feature search is slowed when consecutive targets are defined in different visual dimensions. Behavioral data provide evidence that attentional weight needs to be shifted between dimension-specific processing modules. We found similar dimension-specific change effects in a conjunction search task, in which observers searched for an odd-one-out target defined by a unique combination of size and color or, respectively, size and motion direction. Changes of the secondary target dimension (color or motion) across trials, but not target feature changes within a dimension, increased the time required to detect the target. Dimensional change costs were greatly increased for singleton conjunction search compared to singleton feature search. This suggests involvement of top-down control processes in dimensional change in conjunction search, in contrast to stimulus-driven dimensional change in singleton feature search. The functional anatomical correlates of top-down controlled visual dimension changes were investigated in two event-related functional magnetic resonance imaging (fMRI) experiments. In Experiment 1, dimensional change in singleton conjunction search was accompanied by transient activations in a fronto-posterior network of brain areas that was largely non-overlapping with the general network activated during visual search. Experiment 2, which contrasted singleton feature and conjunction search within the same session, revealed a double dissociation in anterior prefrontal cortex: left frontopolar cortex was selectively involved in stimulus-driven dimension changes but not in top-down controlled dimension changes, whereas the reverse was observed in frontomedian cortex.     

Role of the medial part of the intraparietal sulcus in implementing movement direction

The contribution of the posterior parietal cortex (PPC) to visually guided movements has been originally inferred from observations made in patients suffering from optic ataxia. Subsequent electrophysiological studies in monkeys and functional imaging data in humans have corroborated the key role played by the PPC in sensorimotor transformations underlying goal-directed movements, although the exact contribution of this structure remains debated. Here, we used transcranial magnetic stimulation (TMS) to interfere transiently with the function of the left or right medial part of the intraparietal sulcus (mIPS) in healthy volunteers performing visually guided movements with the right hand. We found that a "virtual lesion" of either mIPS increased the scattering in initial movement direction (DIR), leading to longer trajectory and prolonged movement time, but only when TMS was delivered 100-160 ms before movement onset and for movements directed toward contralateral targets. Control experiments showed that deficits in DIR consequent to mIPS virtual lesions resulted from an inappropriate implementation of the motor command underlying the forthcoming movement and not from an inaccurate computation of the target localization. The present study indicates that mIPS plays a causal role in implementing specifically the direction vector of visually guided movements toward objects situated in the contralateral hemifield.     

A discrete area within the left dorsolateral prefrontal cortex involved in visual-verbal incongruence judgment

The role of the frontal lobe in cross-modal visual-auditory processing has been documented in experiments using incongruent/congruent paradigms. In this study, 4 patients with left frontal World Health Organization Grade II glioma were assessed during pre-, intra-, and postoperative sessions with picture-naming and verbal-visual task requiring judgment of congruence between pictures and words. During awake brain surgery, the naming and cross-modal tasks were coupled with electrical stimulation inactivating restricted specific regions. For all patients, focal brain stimulation in the dorsolateral prefrontal cortex elicited picture-word matching disturbances but no naming impairment, and the elicited errors exclusively appeared in incongruent and not congruent conditions. The dissociation observed between correct picture naming and disturbed cross-modal judgment shows that electrical stimulation of a discrete cortical area within the left dorsolateral prefrontal cortex can inhibit the simultaneous processing of visual-verbal information without disturbing larger networks involved in the naming process.     

Right parietal cortex plays a critical role in change blindness

There is increasing evidence from functional magnetic resonance imaging (fMRI) that visual awareness is not only associated with activity in ventral visual cortex but also with activity in the parietal cortex. However, due to the correlational nature of neuroimaging, it remains unclear whether this parietal activity plays a causal role in awareness. In the experiment presented here we disrupted activity in right or left parietal cortex by applying repetitive transcranial magnetic stimulation (rTMS) over these areas while subjects attempted to detect changes between two images separated by a brief interval (i.e. 1-shot change detection task). We found that rTMS applied over right parietal cortex but not left parietal cortex resulted in longer latencies to detect changes and a greater rate of change blindness compared with no TMS. These results suggest that the right parietal cortex plays a critical role in conscious change detection.     

Role of prefrontal and anterior cingulate regions in decision-making processes shared by memory and nonmemory tasks

In the episodic retrieval (ER) domain, activations in right dorsolateral prefrontal cortex (DLPFC) are often attributed to postretrieval monitoring. Yet, right DLPFC activations are also frequently found during nonmemory tasks. To investigate the role of this region across different cognitive functions, we directly compared brain activity during ER and visual perception (VP) using event-related functional magnetic resonance imaging. In the ER task, participants decided whether words were old or new, whereas in the VP task, they decided which of the two colored screen areas was larger. In both tasks, each decision was followed by a confidence rating. The main finding was that right DLPFC (Brodmann area 46/10) activity was greater for low- than for high-confidence decisions in both tasks, demonstrating a general role in decision making. Even when reaction times (RTs) were included in the model, confidence remained the significant predictor of activity, suggesting that right DLPFC is involved in discontinuous evaluation rather than in continuous monitoring. In contrast, activity in anterior cingulate cortex was not only greater for low-confidence decisions but also increased with RT, reflecting a role in continuous conflict monitoring. Overall, the results demonstrate how direct cross-function comparisons clarify the generality and specificity of the functions of various brain regions.     

A comparison of visual and auditory motion processing in human cerebral cortex

Visual and auditory motion information can be used together to provide complementary information about the movement of objects. To investigate the neural substrates of such cross-modal integration, functional magnetic resonance imaging was used to assess brain activation while subjects performed separate visual and auditory motion discrimination tasks. Areas of unimodal activation included the primary and/or early sensory cortex for each modality plus additional sites extending toward parietal cortex. Areas conjointly activated by both tasks included lateral parietal cortex, lateral frontal cortex, anterior midline and anterior insular cortex. The parietal site encompassed distinct, but partially overlapping, zones of activation in or near the intraparietal sulcus (IPS). A subsequent task requiring an explicit cross-modal speed comparison revealed several foci of enhanced activity relative to the unimodal tasks. These included the IPS, anterior midline, and anterior insula but not frontal cortex. During the unimodal auditory motion task, portions of the dorsal visual motion system showed signals depressed below resting baseline. Thus, interactions between the two systems involved either enhancement or suppression depending on the stimuli present and the nature of the perceptual task. Together, these results identify human cortical regions involved in polysensory integration and the attentional selection of cross-modal motion information.