Search

Cerebellar patients have been found to show deficits in visual motion discrimination, suggesting that the cerebellum may play a role in visual sensory processing beyond mediating motor control. Here we show that triple-pulse online transcranial magnetic stimulation (TMS) over cerebellar vermis but not over the cerebellar hemispheres significantly impaired motion discrimination. Critically, the interference caused by vermis TMS on motion discrimination did not depend on an indirect effect of TMS over nearby visual areas, as demonstrated by a control experiment in which TMS over V1 but not over cerebellar vermis significantly impaired orientation discrimination. These findings demonstrate the causal role of the cerebellar vermis in visual motion processing in neurologically normal participants., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

In this study, we investigated whether early deafness affects the typical pattern of hemispheric lateralization [i.e., right hemisphere (RH) dominance] in the control of spatial attention. To this aim, deaf signers, deaf non-signers, hearing signers, and hearing non-signers were required to bisect a series of centrally presented visual lines. The directional bisection bias was found to be significantly different between hearing and deaf participants, irrespective of sign language use. Hearing participants (both signers and non-signers) showed a consistent leftward bias, reflecting RH dominance. Conversely, we observed no evidence of a clear directional bias in deaf signers or non-signers (deaf participants overall showing a non-significant tendency to deviate rightward), suggesting that deafness may be associated to a more bilateral hemispheric engagement in visuospatial tasks.

Consistent evidence suggests that pitch height may be represented in a spatial format, having both a vertical and a horizontal representation. The spatial representation of pitch height results into response compatibility effects for which high pitch tones are preferentially associated to up-right responses, and low pitch tones are preferentially associated to down-left responses (i.e., the Spatial-Musical Association of Response Codes (SMARC) effect), with the strength of these associations depending on individuals' musical skills. In this study we investigated whether listening to tones of different pitch affects the representation of external space, as assessed in a visual and haptic line bisection paradigm, in musicians and non musicians. Low and high pitch tones affected the bisection performance in musicians differently, both when pitch was relevant and irrelevant for the task, and in both the visual and the haptic modality. No effect of pitch height was observed on the bisection performance of non musicians. Moreover, our data also show that musicians present a (supramodal) rightward bisection bias in both the visual and the haptic modality, extending previous findings limited to the visual modality, and consistent with the idea that intense practice with musical notation and bimanual instrument training affects hemispheric lateralization.

A balance of mutual tonic inhibition between bi-hemispheric posterior parietal cortices is believed to play an important role in bilateral visual attention. However, experimental support for this notion has been mainly drawn from clinical models of unilateral damage. We have previously shown that low-frequency repetitive TMS (rTMS) over the intraparietal sulcus (IPS) generates a contralateral attentional deficit in bilateral visual tracking. Here, we used functional magnetic resonance imaging (fMRI) to study whether rTMS temporarily disrupts the inter-hemispheric balance between bilateral IPS in visual attention. Following application of 1 Hz rTMS over the left IPS, subjects performed a bilateral visual tracking task while their brain activity was recorded using fMRI. Behaviorally, tracking accuracy was reduced immediately following rTMS. Areas ventro-lateral to left IPS, including inferior parietal lobule (IPL), lateral IPS (LIPS), and middle occipital gyrus (MoG), showed decreased activity following rTMS, while dorsomedial areas, such as Superior Parietal Lobule (SPL), Superior occipital gyrus (SoG), and lingual gyrus, as well as middle temporal areas (MT+), showed higher activity. The brain activity of the homologues of these regions in the un-stimulated, right hemisphere was reversed. Interestingly, the evolution of network-wide activation related to attentional behavior following rTMS showed that activation of most occipital synergists adaptively compensated for contralateral and ipsilateral decrement after rTMS, while activation of parietal synergists, and SoG remained competing. This pattern of ipsilateral and contralateral activations empirically supports the hypothesized loss of inter-hemispheric balance that underlies clinical manifestation of visual attentional extinction.

The human capacity to discriminate among different faces relies on distinct parallel subprocesses, based either on the analysis of configural aspects or on the sequential analysis of the single elements of a face. A particular type of configural processing consists of considering whether two faces differ in terms of internal spacing among their features, referred to as second-order relations processing. Findings from electrophysiological, neuroimaging, and lesion studies suggest that, overall, configural processes rely more on the right hemisphere, whereas analysis of single features would involve more the left. However, results are not always consistent, and behavioral evidence for a right-hemisphere specialization in second-order relations processing is lacking. Here, we used divided visual field presentation to investigate the possible different contributions of the two hemispheres to face discrimination based on relational versus featural processing. Our data indicate a right-hemispheric specialization in relational processing of upright (but not inverted) faces. Furthermore, we provide evidence regarding the involvement of both the right and left hemispheres in the processing of faces differing for inner features, suggesting that both analytical and configural modes of processing are at play.

The suggestion has recently been made that certain higher-order cortical areas involved in supporting multisensory representations of the body, and of the space around it, might also play a role in controlling thermoregulatory functions. Here we demonstrate that temporary interference with the function of one of these areas, the posterior parietal cortex, by repetitive transcranial magnetic stimulation, results in a decrease in limb temperature. By contrast, interference with the activity of a sensory-specific area (the primary somatosensory cortex) had no effect on temperature. The results of this experiment suggest that associative multisensory brain areas might exert a top-down modulation over basic physiological control. Such a function might be part of a larger neural circuit responsible for maintaining the integrity of the body at both a homeostatic and a psychological level.

Despite the fact that bilateral mirror symmetry is an important characteristic of the visual world, few studies have investigated its neural basis. Here we addressed this issue by investigating whether the object-selective lateral occipital (LO) cortex, a key brain region in object and shape processing, is causally involved in bilateral symmetry detection. Participants were asked to discriminate between symmetric and asymmetric dot patterns, while fMRI-guided repetitive TMS was delivered online over either the left LO, the right LO or two control sites in the occipital cortex. We found that the application of TMS over both right and left LO impaired symmetry judgments, with disruption being greater following right LO than left LO TMS, indicative of right hemisphere lateralization in symmetry processing. TMS over LO bilaterally also affected a visual contour detection task, with no evidence for hemispheric difference in this task. Overall, our results demonstrates that LO bilaterally plays a causal role in symmetry detection possibly due to symmetry acting as a strong cue in Gestalt processes mediating object recognition., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

Research on visual attention triggered by face gender is still relatively sparse. In the present study, three experiments are reported in which male and female participants were required to estimate the midpoint of a line (i.e., the "line bisection task"): at each end of the line a face was presented. Depending on the experimental condition, faces could be of the same gender (i.e., two males or two females) or the opposite gender. Experiments 1 and 2 converged in showing that when a male face was presented at the right and a female face at the left endpoint of the line, a clear rightward bias emerged compared to the other experimental conditions, indicating that male faces captured attention more than female faces. Importantly, male faces used across Experiments 1 and 2 were rated as more threatening than female faces, suggesting that perceived level of threat may have been responsible for the observed bias toward the male face. Experiment 3 corroborated this hypothesis by finding an attentional bias toward the male face with high threat (angry) faces but not with low threat (smiling) faces.

Emotion recognition is mediated by a complex network of cortical and subcortical areas, with the two hemispheres likely being differently involved in processing positive and negative emotions. As results on valence-dependent hemispheric specialisation are quite inconsistent, we carried out three experiments with emotional stimuli with a task being sensitive to measure specific hemispheric processing. Participants were required to bisect visual lines that were delimited by emotional face flankers, or to haptically bisect rods while concurrently listening to emotional vocal expressions. We found that prolonged (but not transient) exposition to concurrent happy stimuli significantly shifted the bisection bias to the right compared to both sad and neutral stimuli, indexing a greater involvement of the left hemisphere in processing of positively connoted stimuli. No differences between sad and neutral stimuli were observed across the experiments. In sum, our data provide consistent evidence in favour of a greater involvement of the left hemisphere in processing positive emotions and suggest that (prolonged) exposure to stimuli expressing happiness significantly affects allocation of (spatial) attentional resources, regardless of the sensory (visual/auditory) modality in which the emotion is perceived and space is explored (visual/haptic).

Facial recognition relies on distinct and parallel types of processing: featural processing focuses on the individual components of a face (e.g., the shape or the size of the eyes), whereas configural (or "relational") processing considers the spatial interrelationships among the single facial components (e.g., distance of the mouth from the nose). Previous neuroimaging evidence has suggested that featural and configural processes may rely on different brain circuits. By using rTMS, here we show for the first time a double dissociation in dorsolateral prefrontal cortex for different aspects of face processing: in particular, TMS over the left middle frontal gyrus (BA8) selectively disrupted featural processing, whereas TMS over the right inferior frontal gyrus (BA44) selectively interfered with configural processing of faces. By establishing a causal link between activation in left and right prefrontal areas and different modes of face processing, our data extend previous neuroimaging evidence and may have important implications in the study of face-processing deficits, such as those manifested in prosopagnosia and autistic spectrum disorders., (Copyright © 2013 Elsevier Inc. All rights reserved.)

This study assessed whether numerical magnitude affects the setting of basic spatial coordinates and reference frames, namely the subjective straight ahead. Three tasks were given to 24 right-handed healthy participants: a proprioceptive and a visuo-proprioceptive task, requiring pointing to the subjective straight ahead, and a visual task, requiring a perceptual judgment about the straight ahead position of a light moving left-to-right, or right-to-left. A control task, requiring the bisection of rods of different lengths, was also given. The four tasks were performed under conditions of passive auditory numerical (i.e., listening to small, "2," and large, "8," numbers), and neutral auditory-verbal ("blah") stimulation. Numerical magnitude modulated the participants' deviations in the visual straight ahead task, when the movement of the light was from left-to-right, with the small number bringing about a leftward deviation, the large number a rightward deviation. A similar directional modulation was found in the rod bisection task, in line with previous evidence. No effects of numerical magnitude were found on the proprioceptive and visuo-proprioceptive straight ahead tasks. These results suggest that the spatial effects induced by the activation of the mental number line extend to an egocentric frame of reference but only when a portion of horizontal space has to be "actively" explored.

The ability to identify faces is of critical importance for normal social interactions. Previous evidence suggests that early visual deprivation may impair certain aspects of face recognition. The effects of strabismic amblyopia on face processing have not been investigated previously. In this study, a group of individuals with amblyopia were administered two tasks known to selectively measure face detection based on a Gestalt representation of a face (Mooney faces task) and featural and relational processing of faces (Jane faces task). Our data show that--when relying on their amblyopic eye only - strabismic amblyopes perform as well as normally sighted individuals in face detection and recognition on the basis of their single features. However, they are significantly impaired in discriminating among different faces on the basis of the spacing of their single features (i.e., configural processing of relational information). Our findings are the first to demonstrate that strabismic amblyopia may cause specific deficits in face recognition, and add to previous reports characterizing visual perceptual deficits associated in amblyopia as high-level and not only as low-level processing., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

The relationship between the objective accuracy of visual short-term memory (VSTM) representations and their subjective conscious experience is unknown. We investigated this issue by assessing how the objective and subjective components of VSTM in a delayed cue-target orientation discrimination task are affected by intervening distracters. On each trial, participants were shown a memory cue (a grating), the orientation of which they were asked to hold in memory. On approximately half of the trials, a distracter grating appeared during the maintenance interval; its orientation was either identical to that of the memory cue, or it differed by 10° or 40°. The distracters were masked and presented briefly, so they were only consciously perceived on a subset of trials. At the end of the delay period, a memory test probe was presented, and participants were asked to indicate whether it was tilted to the left or right relative to the memory cue (VSTM accuracy; objective performance). In order to assess subjective metacognition, participants were asked indicate the vividness of their memory for the original memory cue. Finally, participants were asked rate their awareness of the distracter. Results showed that objective VSTM performance was impaired by distracters only when the distracters were very different from the cue, and that this occurred with both subjectively visible and invisible distracters. Subjective metacognition, however, was impaired by distracters of all orientations, but only when these distracters were subjectively invisible. Our results thus indicate that the objective and subjective components of VSTM are to some extent dissociable.

Visual stimuli that exhibit vertical symmetry are easier to remember than stimuli symmetric along other axes, an advantage that extends to the haptic modality as well. Critically, the vertical symmetry memory advantage has not been found in early blind individuals, despite their overall superior memory, as compared with sighted individuals, and the presence of an overall advantage for identifying symmetric over asymmetric patterns. The absence of the vertical axis memory advantage in the early blind may depend on their total lack of visual experience or on the effect of prolonged visual deprivation. To disentangle this issue, in this study, we measured the ability of late blind individuals to remember tactile spatial patterns that were either vertically or horizontally symmetric or asymmetric. Late blind participants showed better memory performance for symmetric patterns. An additional advantage for the vertical axis of symmetry over the horizontal one was reported, but only for patterns presented in the frontal plane. In the horizontal plane, no difference was observed between vertical and horizontal symmetric patterns, due to the latter being recalled particularly well. These results are discussed in terms of the influence of the spatial reference frame adopted during exploration. Overall, our data suggest that prior visual experience is sufficient to drive the vertical symmetry memory advantage, at least when an external reference frame based on geocentric cues (i.e., gravity) is adopted.

Patients affected by Parkinson's disease (PD) can provide crucial information about the involvement of the motor system and prefrontal cortex in processing idioms including action verbs, since dopamine modulates the activity of these structures, and, consequently, different levels of this neurotransmitter can induce different cognitive impairments. In order to investigate the ability to process ambiguous idioms containing an action verb in patients, we asked 15 PD patients, in both OFF- and ON-phases, and 15 healthy matched participants to judge the plausibility of literal and idiomatic sentences, each presented at a self-paced rate. Patients in OFF-phase were faster in reading idiomatic than literal sentences, supporting the view that the motor system is not involved in online idiom processing. However, patients during OFF-phase were impaired in judging the plausibility of idiomatic ambiguous sentences, possibly due to the reduction of dopamine in prefrontal regions. The involvement of the motor system was evident in the ON-phase for literal sentences, suggesting that motor activation is strictly dependent on the context.

In two experiments, we combined a semantic blocked naming paradigm with anodal transcranial direct current stimulation (tDCS) to shed light on the neural correlates of the semantic interference (SI) effect. In particular, prior to the naming task, anodal tDCS was applied over the left superior temporal gyrus (STG, Experiment 1) or the left inferior frontal gyrus (IFG, Experiment 2) to enhance cortical excitability in these regions. In both experiments, participants were tested in two sessions in which either real or sham tDCS was delivered. We found that anodal tDCS over the left STG significantly increased the SI effect, whereas anodal tDCS over the left IFG led to a reduction of the SI effect. Overall, our data confirm the existence of a distributed cortical network involved in lexical retrieval and show that both the left IFG and the left STG play a causal role in this process. In particular, the left IFG is likely to be critical in resolving the conflict between competitor lexical representations, while the left STG seems to be the neural locus of the lexical representational system, where competition among different lexical representations occurs., (Copyright © 2012 IBRO. Published by Elsevier Ltd. All rights reserved.)

There is evidence that humans represent numbers in the form of a mental number line (MNL). Here we show that the MNL modulates the representation of visual and haptic space both in healthy individuals and right-brain-damaged patients, both with and without left unilateral spatial neglect (USN). Participants were asked to estimate the midpoint of visually or haptically explored rods while listening to task-irrelevant stimuli: a small digit ("2"), a large digit ("8"), or a non-numerical auditory stimulus ("blah"). In a control silent condition, the bisection error of USN patients was biased rightwards (namely, the marker of USN) only in the visual modality. Regardless of the direction of the bisection error committed in silent trials, listening to the small digit shifted the perceived midline leftwards, and listening to the large digit shifted the perceived midline rightwards, compared to a control condition in which a neutral syllable ("blah") was presented. The shift induced by listening to numbers occurred independently of the modality of response (i.e., both in vision and haptics), and in every group of participants. Interestingly, the effect of auditory numbers processing on space estimation was overall larger for haptically than for visually explored space in all participants. In conclusion, the present data show that listening to irrelevant numbers affects space perception also in patients with left USN, indicating that the spatial representation and attention processes disrupted by USN are not involved in these numerical magnitude-spatial effects., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

The extent to which the generation of mental images draws on the neuronal representations involved in visual perception has been the subject of much debate. To investigate this overlap, we assessed whether adaptation to visual stimuli affects the ability to generate visual mental images; such cross-adaptation would indicate shared neural representations between visual perception and imagery. Mental imagery was tested using a modified version of the clock task, in which subjects are presented with a digital time (e.g. "2.15") and are asked to generate a mental image of the clock hands displaying this time on an empty clock face. Participants were adapted to oriented lines either on the upper or lower side of the clock face prior to the mental image generation. The results showed that mental imagery was impaired when the mental image had to be generated in the adapted region of visual space (Experiment 1). In Experiment 2, we used TMS to determine whether this adaptation effect occurs in the early visual cortex (EVC; V1/V2). Relative to control conditions (No TMS and Vertex TMS), EVC TMS facilitated mental imagery generation when the mental image spatially overlapped with the adapter. Our results thus show that neuronal representations in the EVC which encode (and are suppressed by) visual input play a causal role in visual mental imagery., (Copyright © 2011 Elsevier Inc. All rights reserved.)

Neurologically intact individuals usually show a leftward bias in line bisection, a tendency known as "pseudoneglect", likely reflecting a right-hemisphere dominance in controlling the allocation of spatial attention. Studies in brain-damaged patients with left visuospatial neglect have reported that auditory stimulation may reduce the deficit, both in a spatially dependent and in a spatially independent way. Here we show for the first time that the concurrent binaural presentation of auditory white noise affects healthy individuals' performance in both visual and haptic bisection, reducing their leftward error. We suggest that this effect depends on the noise boosting alertness and restoring the hemispheric activation balance. Our data clearly show that task-irrelevant auditory noise crossmodally affects the allocation of spatial resources in both the haptic and the visual space; future research may clarify whether these effects are specific for the type of auditory stimulation.

The human visual system is able to efficiently extract symmetry information from the visual environment. Prior neuroimaging evidence has revealed symmetry-preferring neuronal representations in the dorsolateral extrastriate visual cortex; the objective of the present study was to investigate the necessity of these representations in symmetry discrimination. This was accomplished by the use of state-dependent transcranial magnetic stimulation, which combines the fine resolution of adaptation paradigms with the assessment of causality. Subjects were presented with adapters and targets consisting of dot configurations that could be symmetric along either the vertical or horizontal axis (or they could be non-symmetric), and they were asked to perform a symmetry discrimination task on the targets while fixating the center of the screen. TMS was applied during the delay between the adapter and the test stimulus over one of four different sites: Left or Right V1/V2, or left or right dorsolateral extrastriate cortex (DLO). TMS over both Left and Right DLO reduced the adaptation effect in detecting vertical and horizontal symmetry, although the Left DLO effect on horizontal symmetry and the Right DLO effect on both vertical and horizontal symmetry were present only when considering subjects who showed a behavioral adaptation effect in the baseline No-TMS condition. Application of TMS over the Left or Right V1/V2 did not modulate the adaptation effect. Overall, these data suggest that both the Left and Right DLO contain neuronal representations tuned to mirror symmetry which play a causal role in symmetry discrimination., (Copyright © 2011 Elsevier Inc. All rights reserved.)

Our representation of peripersonal space does not always accurately reflect the physical world. An example of this is pseudoneglect, a phenomenon in which neurologically normal individuals bisect to the left of the veridical midpoint, reflecting an overrepresentation of the left portion of space compared with the right one. Consistent biases have also been observed in the vertical and radial planes. It is an open question whether these biases depend on normal visual experience for their occurrence. Here we systematically investigated this issue by testing blindfolded sighted and early blind individuals in a haptic line bisection task. Critically, we found a robust leftward bias in all participants. In the vertical and radial planes, sighted participants showed a consistent downward and proximal bias. Conversely, the directional bias in blind participants was dependent on the final movement direction; thus, there was no general bias in either direction. These findings are discussed in terms of different reference frames adopted by sighted and blind participants when encoding space.

Previous studies have demonstrated that transcranial direct current stimulation (tDCS) can be proficiently used to modulate attentional and cognitive functions. For instance, in the language domain there is evidence that tDCS can fasten picture naming in both healthy individuals and aphasic patients, or improve grammar learning. In this study, we investigated whether tDCS can be used to increase healthy subjects' performance in phonemic and semantic fluency tasks, that are typically used in clinical assessment of language. Ten healthy individuals performed a semantic and a phonemic fluency task following anodal tDCS applied over Broca's region. Each participant underwent a real and a sham tDCS session. Participants were found to produce more words following real anodal tDCS both in the phonemic and in the semantic fluency. Control experiments ascertained that this finding did not depend upon unspecific effects of tDCS over levels of general arousal or attention or upon participants' expectations. These data confirm the efficacy of tDCS in transiently improving language functions by showing that anodal stimulation of Broca's region can enhance verbal fluency. Implications of these results for the treatment of language functions in aphasia are considered., (Copyright © 2011 IBRO. Published by Elsevier Ltd. All rights reserved.)

Stereotypes associated with gender, race, ethnicity and religion are powerful forces in human social interactions. Previous neuroimaging and neuropsychological studies point to a role of the prefrontal cortex in controlling stereotypical responses. Here we used transcranial magnetic stimulation (TMS) in combination with an Implicit Association Test (IAT) to highlight the possible causal role of the left dorsolateral prefrontal cortex (DLPFC) and the right anterior dorsomedial prefrontal cortex (aDMPFC) in controlling gender-stereotypical responses. Young male and female participants were tested. Our results showed that applying TMS over the left DLPFC and the right aDMPFC increased the gender-stereotypical bias in male participants compared to when TMS was applied to a control site (vertex). This suggests that both the left DLPFC and the right aDMPFC play a direct role in stereotyping. Females did not show a significant gender bias on the IAT; correspondingly their responses were unaffected by TMS., (Copyright © 2011 Elsevier Inc. All rights reserved.)

Neurologically normal individuals typically show a leftward bias--known as pseudoneglect--in bisecting physical lines as well as numerical intervals. The latter bias may reflect the spatial nature in which numbers are represented (i.e., the mental number line). In previous studies, we found that congenitally blind individuals show such leftward bias in haptic bisection. Here, we demonstrate that blind individuals also show a consistent leftward bias in numerical bisection. The leftward bias was greater when numbers were presented in descending rather than ascending order, and the magnitude of the leftward bias was comparable to that shown by a control group of blindfolded sighted participants. Our findings thus support the view that pseudoneglect operates at a mental representational level rather than being perceptually based. Moreover, the consistent leftward bias shown by blind individuals in both line and numerical bisection suggests that the right hemisphere dominance in spatial processing, resulting in an overestimation of the left side of space, develops even in the absence of any visual input.

The medial prefrontal cortex (mPFC) and the right somatosensory cortex (rSC) are known to be involved in emotion processing and face expression recognition, although the possibility of segregated circuits for specific emotions in these regions remains unclear. To investigate this issue, we used transcranial magnetic stimulation (TMS) together with a priming paradigm to modulate the activation state of the mPFC and the rSC during emotional expressions discrimination. This novel paradigm allows analyzing how TMS interacts with the ongoing activity of different neuronal populations following prime processing. Participants were asked to discriminate between angry and happy faces that were preceded by a congruent prime (a word expressing the same emotion), an incongruent prime (a word expressing the opposite emotion) or a neutral prime. In TMS trials, a single pulse was delivered over the mPFC, rSC or Vertex (control site) between prime and target presentation. TMS applied over the mPFC significantly affected the priming effect, by selectively increasing response latencies in congruent trials. This indicates that the mPFC contains different neural representations for angry and happy expressions. TMS over rSC did not significantly affect the priming effect, suggesting that rSC is not involved in processing verbal emotional stimuli., (Copyright © 2011 Elsevier Ltd. All rights reserved.)

The human posterior parietal cortex (PPC) is involved in the encoding of both visual motion and numerical magnitude. In non human primates, neurons have been found in PPC that are selective for both motion direction and magnitude. Whether such neurons also exist in human PPC is not known. Here we investigated this hypothesis using state-dependent transcranial magnetic stimulation (TMS). Participants were adapted to a specific motion direction (either leftward or rightward), after which they performed a magnitude comparison task, with TMS applied at the onset of each trial. Our hypothesis was that neurons tuned to leftward motion may also be sensitive to small magnitudes and neurons tuned to rightward motion may also be sensitive to large magnitudes, a mapping that may have developed via spatial attentional mechanisms. Our results supported this view by showing that the effect of PPC TMS on small and large numbers depended on the motion direction being adapted, thus suggesting that there may be a functional overlap in neuronal representations of motion direction and numerical magnitude in human PPC., (Copyright © 2011 Elsevier Ireland Ltd. All rights reserved.)

Although much is known about the impact of stimulus properties such as luminance contrast, spatial frequency, and orientation on visually evoked neural activity, much less is known about how they modulate neural activity when they are properties of a mental image held in working memory (WM). Here we addressed this question by investigating how a parametric manipulation of an imagined stimulus attribute affects neuronal excitability in the early visual cortex. We manipulated luminance contrast, a stimulus property known to strongly affect the magnitude of neuronal responses in early visual areas. Luminance contrast modulated neuronal excitability, as assessed by the frequency of phosphenes induced by transcranial magnetic stimulation (TMS) with the exact nature of this modulation depending on TMS intensity. These results point to a strong overlap in the neuronal processes underlying visual perception and mental imagery: not only does WM maintenance selectively engage neurons which are tuned to the maintained attribute (as has previously been shown), but the extent to which those neurons are activated depends on the image contrast (as is the case with visually evoked responses). From a methodological viewpoint, these results suggest that assessment of visual cortical excitability using TMS is affected by the TMS intensity used to probe the neuronal population.

Numerical magnitude is believed to be represented along a mental number line (MNL), and there is evidence to suggest that the activation of the MNL affects the perception and representation of external space. In the present study, we investigated whether a spatial motor task affects numerical processing in the auditory modality. Blindfolded participants were presented with a numerical interval bisection task, while performing a tapping task with either their left or right hand, either in the fronto-central, fronto-left, or fronto-right peripersonal space. Results showed that tapping significantly influenced the participants' numerical bisection, with tapping in the left side of space increasing the original tendency to err leftward, and tapping to the right reducing such bias. Importantly, the effect depended on the side of space in which the tapping activity was performed, regardless of which hand was used. Tapping with either the left or right hand in the fronto-central space did not affect the participants' bias. These findings offer novel support for the existence of bidirectional interactions between external and internal representations of space.

Bilateral mirror symmetry, especially vertical symmetry, is a powerful phenomenon in spatial organization of visual shapes. However, the causes of vertical symmetry salience in visual perception are not completely clear. Here we investigated whether the perceptual salience of vertical symmetry depends on visual experience by testing a group of congenitally blind individuals in a memory task in which either horizontal or vertical symmetry was used as an incidental feature. Both blind and sighted subjects remembered more accurately configurations that were symmetrical compared to those that were not. Critically, whereas sighted subjects displayed a higher level of facilitation by vertical than horizontal symmetry, no such difference was found in the blind. This suggests that the perceptual salience of the vertical dimension is visually based., (2010 Elsevier B.V. All rights reserved.)

Cortical areas involved in sensory analysis are also believed to be involved in short-term storage of that sensory information. Here we investigated whether transcranial magnetic stimulation (TMS) can reveal the content of visual short-term memory (VSTM) by bringing this information to visual awareness. Subjects were presented with two random-dot displays (moving either to the left or to the right) and they were required to maintain one of these in VSTM. In Experiment 1, TMS was applied over the motion-selective area V5/MT+ above phosphene threshold during the maintenance phase. The reported phosphene contained motion features of the memory item, when the phosphene spatially overlapped with memory item. Specifically, phosphene motion was enhanced when the memory item moved in the same direction as the subjects' V5/MT+ baseline phosphene, whereas it was reduced when the motion direction of the memory item was incongruent with that of the baseline V5/MT+ phosphene. There was no effect on phosphene reports when there was no spatial overlap between the phosphene and the memory item. In Experiment 2, VSTM maintenance did not influence the appearance of phosphenes induced from the lateral occipital region. These interactions between VSTM maintenance and phosphene appearance demonstrate that activity in V5/MT+ reflects the motion qualities of items maintained in VSTM. Furthermore, these results also demonstrate that information in VSTM can modulate the pattern of visual activation reaching awareness, providing evidence for the view that overlapping neuronal populations are involved in conscious visual perception and VSTM., (2010. Published by Elsevier Inc.)

Humans tend to represent numbers in the form of a mental number line. Here we show that the mental number line can modulate the representation of peripersonal haptic space in a crossmodal fashion and that this interaction is not visually mediated. Sighted and early-blind participants were asked to haptically explore rods of different lengths and to indicate midpoints of those rods. During each trial, either a small (2) or a large (8) number was presented in the auditory modality. When no numbers were presented, participants tended to bisect the rods to the left of the actual midpoint, consistent with the notion of pseudoneglect. In both groups, this bias was significantly increased by the presentation of a small number and was significantly reduced by the presentation of a large number. Hence, spatial shifts of attention induced by number processing are not limited to visual space or embodied responses but extend to haptic peripersonal space and occur crossmodally without requiring the activation of a visuospatial representation.

The left ventral premotor cortex (PMv) is preferentially activated by exemplars of tools, suggestive of category specificity in this region. Here we used state-dependent transcranial magnetic stimulation (TMS) to investigate the causal role of such category-specific neuronal representations in the encoding of tool words. Priming to a category name (either "Tool" or "Animal") was used with the objective of modulating the initial activation state of this region prior to application of TMS and the presentation of the target stimulus. When the target word was an exemplar of the "Tool" category, the effects of TMS applied over PMv (but not PMd) interacted with priming history by facilitating reaction times on incongruent trials while not affecting congruent trials. This congruency/TMS interaction implies that the "Tool" and "Animal" primes had a differential effect on the initial activation state of the left PMv and implies that this region is one neural locus of category-specific behavioral priming for the "Tool" category. TMS applied over PMv had no behavioral effect when the target stimulus was an exemplar of the "Animal" category, regardless of whether the target word was congruent or incongruent with the prime. That TMS applied over the left PMv interacted with a priming effect that extended from the category name ("Tool") to exemplars of that category suggests that this region contains neuronal representation associated with a specific semantic category. Our results also demonstrate that the state-dependent effects obtained in the combination of visual priming and TMS are useful in the study of higher-level cognitive functions., (Copyright (c) 2009 Elsevier Inc. All rights reserved.)

The mental number line has been shown to exert an influence on the visuo-spatial allocation of attention, with presentation of numbers from the low and high ends of the mental number line inducing covert shifts of spatial attention to the left and right side of visual space, respectively. In the present study we used transcranial magnetic stimulation (TMS) to investigate whether this attentional modulation influences excitability of the early visual cortex. By using the phosphene threshold as a measure of visual cortical excitability, we show (in 10 subjects) that number priming modulates excitability of the early visual cortex in a topographic fashion: low numbers, associated with left side of space, increase the excitability of the right early visual cortex (the stimulation of which induces phosphenes in the left hemifield) and decrease the excitability of the left early visual cortex (the stimulation of which induces phosphenes in the right hemifield). The opposite pattern of results was observed for high numbers. Our results suggest that the attentional shifts induced by the mental number line are manifested at the earliest cortical stages of visual processing.

Activation of the left posterior parietal cortex (PPC) has been associated with the encoding of letters independent of visual form. Here we used transcranial magnetic stimulation (TMS)-adaptation to investigate whether this abstract letter selectivity plays a causal role in letter processing. Visual adaptation was used to manipulate the initial activation state of neurons tuned to different letters prior to the application of TMS, after which subjects performed a detection task on letters that were presented in a different case from the adapting letter. After adaptation, TMS applied over the left PPC facilitated the detection of the adapted letter, whereas it had no impact on the detection of nonadapted letters. TMS applied over the right PPC had no significant effect on either type of letter. This interaction between adaptation and the effects of left PPC TMS demonstrates that adaptation modulated neural activity in the left PPC and thus demonstrates abstract letter selectivity in this region. Importantly, as the adapted letter and the target letters were presented in different cases, this finding demonstrates that the left PPC plays a causal role in letter processing independent of visual form.

Whether visual imagery and visual short-term memory (STM) share the same neural resources, and the extent to which the early visual cortex (V1/V2) is involved in these processes, has been the subject of much debate. Here, we used transcranial magnetic stimulation (TMS) in two separate experiments to contrast the neural states associated with visual imagery and visual STM in the early visual cortex. In Experiment 1, we investigated V1/V2 activation states at the end of the retention phase in a visual imagery and a visual STM task. V1/V2 TMS facilitated performance in both tasks; the finding that imagery and STM interacted with TMS in the same way suggests that the two processes have similar effects on early visual cortical excitability. In Experiment 2, we investigated V1/V2 activation states at the beginning of the retention phase. V1/V2 TMS impaired performance in the visual STM task, whereas it had no effect on the imagery task. Taken together, our findings show that the late phases of the early visual cortical activation state associated with visual imagery and visual STM are similar; differences between the two processes are apparent in the early phases of the tasks. Our results also suggest that the causal role of the early visual cortex in visual STM includes both the initial translation of the visual input into working memory and the subsequent maintenance of the mental representation. Finally, our findings indicate that visual STM sensory recruitment in working memory might act via excitability modulation of V1/V2 neurons.

Along with meaning and form, words can be described on the basis of their grammatical properties. Grammatical gender is often used to investigate the latter as it is a grammatical property that is independent of meaning. The left inferior frontal gyrus (IFG) has been implicated in the encoding of grammatical gender, but its causal role in this process in neurologically normal observers has not been demonstrated. Here we combined verbal satiation with transcranial magnetic stimulation (TMS) to demonstrate that subpopulations of neurons within Broca's area respond preferentially to different classes of grammatical gender. Subjects were asked to classify Italian nouns into living and nonliving categories; half of these words were of masculine and the other half of feminine grammatical gender. Prior to each test block, a satiation paradigm (a phenomenon in which verbal repetition of a category name leads to a reduced access to that category) was used to modulate the initial state of the representations of either masculine or feminine noun categories. In the No TMS condition, subjects were slower in responding to exemplars to the satiated category relative to exemplars of the nonsatiated category, implying that the neural representations for different classes of grammatical gender are partly dissociable. The application of TMS over Broca's area removed the behavioral impact of verbal (grammatical) satiation, demonstrating the causal role of this region in the encoding of grammatical gender. These results show that the neural representations for different cases of a grammatical property within Broca's area are dissociable.

Mental representation of numbers is believed to be spatial in nature, with small numbers occupying the left and large numbers the right side of a putative mental number line. Consistent with this, presentation of numbers from the low and high ends of the mental number line induces covert shifts of spatial attention to the left and right side of visual space, respectively. However, the effect of the presentation of the middle range (containing numbers below and above the midpoint) of the number line on visual perception has so far not been studied. Here we show in two experiments, using a line bisection task and a simple target detection task, that processing of middle-range numbers affects allocation of visuospatial attention in a similar way as processing of small numbers, with attention shifted to the left side of space. We suggest that this pattern of results arises due to "anchoring" heuristics that participants use in number processing.

We tend to mentally organize numbers along a left-to-right oriented horizontal mental number line, with the smaller numbers occupying the more leftward positions. This mental number line has been shown to exert an influence on the visuospatial allocation of attention, with presentation of numbers from the low and high ends of the mental number line inducing covert shifts of spatial attention to the left and right side of visual space, respectively. However, the neural basis of this modulation is not known. Here we used transcranial magnetic stimulation (TMS) to study the role of the angular gyrus in shifts in visuospatial attention induced by the mental number line. We used a priming paradigm with a line bisection task to assess the bias in spatial allocation of visual attention induced by exposure to either small (16-24) or large (76-84) ends of the mental number line. In the Small Number Prime condition, when attention is presumably biased to the left side of visual space, TMS applied over the right angular gyrus during the delay between the prime and the target line abolished the effect of number priming. In contrast, application of TMS over the left angular gyrus had no significant effect. In the Large Number Prime condition (which shifted attention to the right side of visual space) both left and right TMS over the angular gyrus modulated the effect of number priming. This pattern of results reveals the involvement of the angular gyrus in the interaction between the mental number line and visual spatial attention.

The state-dependency of transcranial magnetic stimulation (TMS) can be used to investigate the neural properties of subregions of the stimulated region. The objective of the present study was to determine whether state-dependency can reveal letter selectivity in the left posterior parietal cortex (PPC), a region known to contain letter-selective neurons. In two experiments, we used visual priming and adaptation to modulate the initial activation state of the left PPC prior to application of TMS. In the priming experiment, TMS was applied over the left PPC during the delay between the prime and the target stimulus on each experimental trial. Left PPC TMS reversed the effects of priming by facilitating the detection of non-primed letters, whereas detection of primed letters was unaffected. As neurons tuned to non-primed letters were less active at the time of TMS application than neurons tuned to the primed letters, this finding demonstrates that TMS preferentially facilitates the detection of attributes encoded by the less active neural populations. A similar facilitation of the less active neural populations was observed when adaptation was used to suppress letter-selective neurons prior to application of TMS. Our study demonstrates that TMS-priming and TMS-adaptation paradigms can reveal letter selectivity in the left PPC and thus be useful in the study of language processes. Our results also show that the state-dependent TMS effects obtained with visual priming are similar to those found with TMS adaptation: in both cases, attributes encoded by the less active neural populations are preferentially facilitated.

To investigate the relationship between visual acuity and cognitive function with aging, we compared low-vision and normally-sighted young and elderly individuals on a spatial working memory (WM) task. The task required subjects to memorise target locations on different matrices after perceiving them visually or haptically. The haptic modality was included as a control to look at the effect of aging on memory without the confounding effect of visual deficit. Overall, age and visual status did not interact to affect WM accuracy, suggesting that age does not exaggerate the effects of visual deprivation. Young participants performed better than the elderly only when the task required more operational processes (i.e., integration of information). Sighted participants outperformed the visually impaired regardless of testing modality suggesting that the effect of the visual deficit is not confined to only the most peripheral levels of information processing. These findings suggest that vision, being the primary sensory modality, tends to shape the general supramodal mechanisms of memory.

The transcranial magnetic stimulation (TMS)-adaptation paradigm, based on the state-dependency of TMS effects, may become a useful tool for differential stimulation of functionally distinct neural populations within the stimulated region. Here we investigated, in the context of motion perception, the time course of state-dependent TMS effects in this paradigm. After adapting to a motion stimulus, subjects were asked to perform a motion direction discrimination task, with TMS applied over the motion selective area V5/MT prior to each experimental trial. Consistent with previous studies, TMS reversed the behavioral effect of adaptation; that is, detection of the adapted direction was enhanced and that of the unadapted direction was impaired. Importantly, this reversal was consistent over the whole block of trials carried out after adaptation: the state-dependent TMS effect was similar in the first and second halves of the post-adaptation discrimination block. This shows that while single-pulse TMS interacts with the effects of adaptation on a trial-by-trial basis to induce state-dependent effects, it does not abolish the effects of adaptation; rather, after each trial, the stimulated region returns to a state of adaptation.

The objective of this review is to examine and evaluate recent findings on cognitive functioning (in particular imagery processes) in individuals with congenital visual impairments, including total blindness, low-vision and monocular vision. As one might expect, the performance of blind individuals in many behaviours and tasks requiring imagery can be inferior to that of sighted subjects; however, surprisingly often this is not the case. Interestingly, there is evidence that the blind often employ different cognitive mechanisms than sighted subjects, suggesting that compensatory mechanisms can overcome the limitations of sight loss. Taken together, these studies suggest that the nature of perceptual input on which we commonly rely strongly affects the organization of our mental processes. We also review recent neuroimaging studies on the neural correlates of sensory perception and mental imagery in visually impaired individuals that have cast light on the plastic functional reorganization mechanisms associated with visual deprivation.

Monocular deprivation has been associated with both specific deficits and enhancements in visual perception and processing. In this study, performance on a visuo-spatial memory task was compared in congenitally monocular individuals and sighted control individuals viewing monocularly (i.e., patched) and binocularly. The task required the individuals to view and memorize a series of target locations on two-dimensional matrices. Overall, congenitally monocular individuals performed worse than sighted individuals (with a specific deficit in simultaneously maintaining distinct spatial representations in memory), indicating that the lack of binocular visual experience affects the way visual information is represented in visuo-spatial memory. No difference was observed between the monocular and binocular viewing control groups, suggesting that early monocular deprivation affects the development of cortical mechanisms mediating visuo-spatial cognition.

The state-dependency approach of transcranial magnetic stimulation (TMS) enables differential stimulation of functionally distinct neural populations within the affected region of cortex. Here we tested the validity of a paradigm based on state-dependency, the TMS-adaptation paradigm, in the context of visual motion perception. Visual adaptation was used to induce an activity imbalance in direction-selective neurons in the visual cortex, after which participants performed a motion direction discrimination task. When TMS was applied over the motion-selective area V5/MT before each experimental trial, the detection of the direction encoded by the adapted neurons was facilitated relative to the direction encoded by the nonadapted neurons. This finding demonstrates, in the domain of visual motion detection, the state-dependency of TMS effects and the validity of the TMS-adaptation paradigm.

In this article, the authors investigated unimodal and cross-modal processes in spatial working memory. A number of locations had to be memorized within visual or haptic matrices according to different experimental conditions known to be critical in accounting for the effects of perception on imagery. Results reveal that some characteristics of the generated mental image remained strictly inherent to the modality in which information was acquired; in general, accuracy was higher when configurations were visually rather than haptically explored (Experiments 1 and 3). Interestingly, the same pattern emerged when the effects of simultaneous versus sequential processing of the stimuli inherent to vision and haptics were isolated from perceptual modality (Experiment 2). Supramodal elements were also identified (Experiment 3) that were specifically associated to the nature of the cognitive processes, regardless of the original characteristics of the sensory information. These data indicate that both unimodal modality-specific and higher order supramodal mechanisms are simultaneously used in spatial processes.

Transcranial magnetic stimulation (TMS) is increasingly used to modify brain activity noninvasively and to study brain-behavior relations. However, results can be variable and the conditions that affect the functional efficacy of TMS remain unclear. Here we show that on-line TMS can either facilitate or suppress perceptual functions depending on the baseline level of activity of the targeted brain region. When TMS was applied over the motion selective region V5/MT during a simple motion-detection task, subjects' motion-detection ability was impaired. Similarly, suppression of V5/MT activity using off-line 1 Hz repetitive TMS (rTMS) disrupted performance in a subsequent motion-detection task. However, paradoxically, on-line V5/MT TMS had a facilitatory effect on motion detection if V5/MT had been suppressed by off-line 1-Hz rTMS prior to the motion-detection task. These results demonstrate that TMS can have an unexpected facilitatory effect on behavior when the targeted neural population is in a suppressed state. Our findings provide further evidence for the view that the effects of TMS are modulated by the initial activation state of the targeted neural population.

Spatial memory is usually better for iconic than for verbal material. Our aim was to assess whether such effect is related to the way iconic and verbal targets are viewed when people have to memorize their locations. Eye movements were recorded while participants memorized the locations of images or words. Images received fewer, but longer, gazes than words. Longer gazes on images might reflect greater attention devoted to images due to their higher sensorial distinctiveness and/or generation with images of an additional phonological code beyond the visual code immediately available. We found that words were scanned mainly from left to right while a more heterogeneous scanning strategy characterized encoding of images. This suggests that iconic configurations tend to be maintained as global integrated representations in which all the item/location pairs are simultaneously present whilst verbal configurations are maintained through more sequential processes.

Similarly to sighted people, individuals congenitally affected by a severe visual impairment can maintain and mentally manipulate spatial information about tactile stimuli [Vecchi, T., Cattaneo, Z., Monegato, M., Pece, A., Cornoldi, C., Pietrini, P., 2006. Why Cyclops could not compete with Ulysses: monocular vision and mental images. NeuroReport 17, 723-726]. The aim of the present study was to assess whether the onset timing of a severe (but not total) sight loss may influence spatial imagery abilities based on haptic input. To this purpose, a group of late severe visually impaired people and a matched group of normally sighted participants (all blindfolded) were presented with an imagery task requiring to memorize and retrieve a number of locations presented on tactile matrices. Results indicate that a severe visual deficit occurring later in life significantly impairs spatial imagery abilities to a greater extent than in the case of congenital blindness, probably as a consequence of a modest development of specific compensatory mechanisms associated with congenital deficits.

The present research demonstrates that the limitations of congenitally blind people in tasks requiring the processing of mental images are specifically related to the absence of binocular vision and not to the absence of vision per se. We contrasted three different groups of participants: sighted; visually impaired, with reduced binocular vision; monocular, with a normal visual acuity although in one eye only. Visually impaired participants (i.e. blurred vision) show a pattern of performance comparable to that of the sighted. In contrast, monocular participants show a similar pattern of performance to congenitally blind individuals despite being able to see perfectly well. These results shed new light on the relationship between perception and imagery and on the characteristics of sequential and simultaneous processes in the human brain.

It has been hypothesized that gender differences in visuo-spatial working memory (VSWM) are larger in tasks requiring active elaboration of the material. In the present study we explored this issue by using an object relocation task, with both verbal and visual stimuli. The involvement of active processes was manipulated through the type of transformation required on the stimulus and through the introduction of different kinds of interference. In the three experiments reported, participants were shown either words or cartoon object icons in different locations and had to relocate them in either the same format or in the opposite one (object icons could be transformed into words and vice versa). Males outperformed females in the most demanding conditions, in which object icons and words were presented together in the encoding phase, and both had to be transformed in the recall phase; or when more demanding interferences were used. Our data suggest that the retention strategy was similar for the two groups and that the gender effect is related to a selective female difficulty associated with the increase in active VSWM processing. These findings further support the hypothesized distinction between the passive and active components of VSWM and illustrate the role that this distinction might play in accounting for individual differences.