Cerebellar contributions to verbal working memory.

There is increasing evidence for a cerebellar role in working memory. Clinical research has shown that working memory impairments after cerebellar damage and neuroimaging studies have revealed task-specific activation in the cerebellum during working memory processing. A lateralisation of cerebellar function within working memory has been proposed with the right hemisphere making the greater contribution to verbal processing and the left hemisphere for visuospatial tasks. We used continuous theta burst stimulation (cTBS) to examine whether differences in post-stimulation performance could be observed based on the cerebellar hemisphere stimulated and the type of data presented. We observed that participants were significantly less accurate on a verbal version of a Sternberg task after stimulation to the right cerebellar hemisphere when compared to left hemisphere stimulation. Performance on a visual Sternberg task was unaffected by stimulation of either hemisphere. We discuss our results in the context of prior studies that have used cerebellar stimulation to investigate working memory and highlight the cerebellar role in phonological encoding.

Better together? The cognitive advantages of synaesthesia for time, numbers, and space.

Synaesthesia for time, numbers, and space (TNS synaesthesia) is thought to have costs and benefits for recalling and manipulating time and number. There are two competing theories about how TNS synaesthesia affects cognition. The "magnitude" account predicts that TNS synaesthesia may affect cardinal magnitude judgements, whereas the "sequence" account suggests that it may affect ordinal sequence judgements and could rely on visuospatial working memory. We aimed to comprehensively assess the cognitive consequences of TNS synaesthesia and distinguish between these two accounts. TNS synaesthetes, grapheme-colour synaesthetes, and nonsynaesthetes completed a behavioural task battery. Three tasks involved cardinal and ordinal comparisons of temporal, numerical, and spatial stimuli; we also examined visuospatial working memory. TNS synaesthetes were significantly more accurate than nonsynaesthetes in making ordinal judgements about space. This difference was explained by significantly higher visuospatial working memory accuracy. Our findings demonstrate an advantage of TNS synaesthesia that is more in line with the sequence account.

Feature integration in visual working memory: parietal gamma activity is related to cognitive coordination.

The mechanism by which distinct subprocesses in the brain are coordinated is a central conundrum of systems neuroscience. The parietal lobe is thought to play a key role in visual feature integration, and oscillatory activity in the gamma frequency range has been associated with perception of coherent objects and other tasks requiring neural coordination. Here, we examined the neural correlates of integrating mental representations in working memory and hypothesized that parietal gamma activity would be related to the success of cognitive coordination. Working memory is a classic example of a cognitive operation that requires the coordinated processing of different types of information and the contribution of multiple cognitive domains. Using magnetoencephalography (MEG), we report parietal activity in the high gamma (80-100 Hz) range during manipulation of visual and spatial information (colors and angles) in working memory. This parietal gamma activity was significantly higher during manipulation of visual-spatial conjunctions compared with single features. Furthermore, gamma activity correlated with successful performance during the conjunction task but not during the component tasks. Cortical gamma activity in parietal cortex may therefore play a role in cognitive coordination.

Strategic resource allocation in the human brain supports cognitive coordination of object and spatial working memory.

The ability to integrate different types of information (e.g., object identity and spatial orientation) and maintain or manipulate them concurrently in working memory (WM) facilitates the flow of ongoing tasks and is essential for normal human cognition. Research shows that object and spatial information is maintained and manipulated in WM via separate pathways in the brain (object/ventral versus spatial/dorsal). How does the human brain coordinate the activity of different specialized systems to conjoin different types of information? Here we used functional magnetic resonance imaging to investigate conjunction- versus single-task manipulation of object (compute average color blend) and spatial (compute intermediate angle) information in WM. Object WM was associated with ventral (inferior frontal gyrus, occipital cortex), and spatial WM with dorsal (parietal cortex, superior frontal, and temporal sulci) regions. Conjoined object/spatial WM resulted in intermediate activity in these specialized areas, but greater activity in different prefrontal and parietal areas. Unique to our study, we found lower temporo-occipital activity and greater deactivation in temporal and medial prefrontal cortices for conjunction- versus single-tasks. Using structural equation modeling, we derived a conjunction-task connectivity model that comprises a frontoparietal network with a bidirectional DLPFC-VLPFC connection, and a direct parietal-extrastriate pathway. We suggest that these activation/deactivation patterns reflect efficient resource allocation throughout the brain and propose a new extended version of the biased competition model of WM.

Neural signatures of stimulus features in visual working memory--a spatiotemporal approach.

We examined the neural signatures of stimulus features in visual working memory (WM) by integrating functional magnetic resonance imaging (fMRI) and event-related potential data recorded during mental manipulation of colors, rotation angles, and color-angle conjunctions. The N200, negative slow wave, and P3b were modulated by the information content of WM, and an fMRI-constrained source model revealed a progression in neural activity from posterior visual areas to higher order areas in the ventral and dorsal processing streams. Color processing was associated with activity in inferior frontal gyrus during encoding and retrieval, whereas angle processing involved right parietal regions during the delay interval. WM for color-angle conjunctions did not involve any additional neural processes. The finding that different patterns of brain activity underlie WM for color and spatial information is consistent with ideas that the ventral/dorsal "what/where" segregation of perceptual processing influences WM organization. The absence of characteristic signatures of conjunction-related brain activity, which was generally intermediate between the 2 single conditions, suggests that conjunction judgments are based on the coordinated activity of these 2 streams.

Electrophysiological correlates of improved short-term memory for emotional faces.

Long-term memory (LTM) is enhanced for emotional information, but the influence of stimulus emotionality on short-term memory (STM) is less clear. We examined the electrophysiological correlates of improved visual STM for emotional face identity, focusing on the P1, N170, P3b and N250r event-related potential (ERP) components. These correlates are taken to indicate which memory processing stages and cognitive processes contribute to the improved STM for emotional face identity. In the encoding phase, one or three angry, happy or neutral faces were presented for 2s, resulting in a memory load of one or three. The subsequent 1-s retention phase was followed by a 2-s retrieval phase, in which participants indicated whether a probe face had been present or not during encoding. Memory performance was superior for angry and happy faces over neutral faces at load three. None of the ERP components during encoding were affected by facial expression. During retrieval, the early P3b was decreased for emotional compared to neutral faces, which presumably reflects greater resource allocation to the maintenance of the emotional faces. Furthermore, the N250r during retrieval was increased for emotional compared to neutral faces, reflecting an enhanced repetition effect for emotional faces. These findings suggest that enhanced visual STM for emotional faces arises from improved maintenance and from improved detection of face repetition at retrieval.

Does transcranial direct current stimulation to prefrontal cortex affect mood and emotional memory retrieval in healthy individuals?

Studies using transcranial direct current stimulation (tDCS) of prefrontal cortex to improve symptoms of depression have had mixed results. We examined whether using tDCS to change the balance of activity between left and right dorsolateral prefrontal cortex (DLPFC) can alter mood and memory retrieval of emotional material in healthy volunteers. Participants memorised emotional images, then tDCS was applied bilaterally to DLPFC while they performed a stimulus-response compatibility task. Participants were then presented with a set of images for memory retrieval. Questionnaires to examine mood and motivational state were administered at the beginning and end of each session. Exploratory data analyses showed that the polarity of tDCS to DLPFC influenced performance on a stimulus-response compatibility task and this effect was dependent on participants' prior motivational state. However, tDCS polarity had no effect on the speed or accuracy of memory retrieval of emotional images and did not influence positive or negative affect. These findings suggest that the balance of activity between left and right DLPFC does not play a critical role in the mood state of healthy individuals. We suggest that the efficacy of prefrontal tDCS depends on the initial activation state of neurons and future work should take this into account.

Cerebellar contributions to spatial memory.

There is mounting evidence for a role for the cerebellum in working memory (WM). The majority of relevant studies has examined verbal WM and has suggested specialisation of the right cerebellar hemisphere for language processing. Our study used theta burst stimulation (TBS) to examine whether there is a converse cerebellar hemispheric specialisation for spatial WM. We conducted two experiments to examine spatial WM performance before and after TBS to mid-hemispheric and lateral locations in the posterior cerebellum. Participants were required to recall the order of presentation of targets on a screen or the targets' order of presentation and their locations. We observed impaired recollection of target order after TBS to the mid left cerebellar hemisphere and reduced response speed after TBS to the left lateral cerebellum. We suggest that these results give evidence of the contributions of the left cerebellar cortex to the encoding and retrieval of spatial information.

Frontal and parietal theta burst TMS impairs working memory for visual-spatial conjunctions.

In tasks that selectively probe visual or spatial working memory (WM) frontal and posterior cortical areas show a segregation, with dorsal areas preferentially involved in spatial (e.g. location) WM and ventral areas in visual (e.g. object identity) WM. In a previous fMRI study [1], we showed that right parietal cortex (PC) was more active during WM for orientation, whereas left inferior frontal gyrus (IFG) was more active during colour WM. During WM for colour-orientation conjunctions, activity in these areas was intermediate to the level of activity for the single task preferred and non-preferred information. To examine whether these specialised areas play a critical role in coordinating visual and spatial WM to perform a conjunction task, we used theta burst transcranial magnetic stimulation (TMS) to induce a functional deficit. Compared to sham stimulation, TMS to right PC or left IFG selectively impaired WM for conjunctions but not single features. This is consistent with findings from visual search paradigms, in which frontal and parietal TMS selectively affects search for conjunctions compared to single features, and with combined TMS and functional imaging work suggesting that parietal and frontal regions are functionally coupled in tasks requiring integration of visual and spatial information. Our results thus elucidate mechanisms by which the brain coordinates spatially segregated processing streams and have implications beyond the field of working memory.

Hemispheric specialisation in haptic processing.

This study investigated the organising principles of touch. We examined specialisations within the haptic system and their hemispheric distribution. Haptic processing consists of the integration of data from multiple sources to form a single percept. Previous research provides strong support for a hierarchical and functional distribution within haptic processing. We investigated hemispheric asymmetry in haptic discrimination of objects with differing textures and centres of mass. By analogy with vision it was hypothesised that participants would demonstrate a left-hand advantage for centre of mass discrimination (a 'global', presumed right hemisphere, judgement) and a right-hand advantage for surface texture judgements (a 'local', presumed left hemisphere discrimination). We found that left-handed participants showed these effects to a lesser degree than did the right-handers, consistent with the notion that left-handed people generally show weaker asymmetries in bimanual tasks. In a second experiment the effect of conflicting information on haptic percept formation was investigated. Following from the previous hypotheses it was predicted that participants would be more accurate with their right hands at judging conflicting surfaces. Contrary to predictions an advantage was demonstrated for the left hand for texture discrimination and for the right hand for centre of mass judgement.

On the functional significance of Novelty-P3: facilitation by unexpected novel sounds.

The unexpected occurrence of a change in the auditory context has been shown to result in distraction due to a momentary attention shift. These unexpected sounds elicit the Novelty-P3 (NP3) response which has been proposed as an electrophysiological index of distractibility, and used as such in the evaluation of several clinical populations. However, unexpected sounds also result in facilitation under certain conditions. Here, we investigate the electrophysiological concomitants of novel sounds in a task in which these sounds facilitate visual task performance. Novel sounds elicited NP3 and resulted in an enhancement of the visual P300 response to subsequent visual targets. This result clearly argues against the use of NP3 as an index of distractibility and asks for a reformulation of the functional significance of this response. We suggest that the NP3 is a complex signal that comprises alerting, orienting and executive control processes triggered by the unexpected stimulus.

Working memory load for faces modulates P300, N170, and N250r.

We used event-related potential (ERP) methodology to examine neural activity associated with visual working memory (WM) for faces. There were two main goals. First, to extend previous findings of P300 load modulation to WM for faces. Second, to examine whether N170 and N250r are also influenced by WM load. Between one and four unfamiliar faces were simultaneously presented for memory encoding. After a 1-sec delay, a target face appeared, and participants had to judge whether this face was part of the previous face array. P300 amplitude decreased as WM load increased, and this P300 suppression was observed at both encoding and retrieval. WM load was also found to modulate other ERPs. The amplitude of the N170 elicited by the target face decreased with load, and this N170 decrease leveled off at load 2, reflecting the behavioral WM capacity of around two faces. In addition, the N250r, observed as an ERP difference for target faces that were present in the encoding array relative to target faces that were absent, was also reduced for higher WM loads. These findings extend previous work by showing that P300 modulation by WM load also occurs for faces. Furthermore, we show, for the first time, that WM load affects the N250r and the early visual N170 component. This suggests that higher visual areas play an important role in WM for faces.

Inhibition of return and action affordances.

Previous work has shown that inhibition of return (IOR) operates on identity-specific object representations and does not spread to semantically related items (Morgan et al. in Eur J Cogn Psychol 17(4):499-520, 2005). The current study further examines the representations underlying IOR by investigating whether IOR is influenced by action-related properties of objects. An exogenous cue appeared on the side of an object that was either compatible or incompatible with the action afforded by the object. Then a target requiring a rapid key-press or reach localisation response appeared on the cued or uncued side. Responses were slower to cued versus uncued targets, but this IOR effect did not interact with action-compatibility. These results show that IOR does not appear to operate on visuomotor representations.