Orienting attention to locations in mental representations.

Many cognitive processes depend on our ability to hold information in mind, often well beyond the offset of the original sensory input. The capacity of this visual short-term memory (VSTM) is limited to around three to four items. Recent research has demonstrated that the content of VSTM can be modulated by top-down attentional biases. This has been demonstrated using retrodictive spatial cues, termed "retro-cues," which orient subjects' attention to spatial locations within VSTM. In the present article, we tested whether the use of these cues is modulated by memory load and cue delay. There are a number of important conclusions: (1) Top-down biases can operate on very brief iconic traces as well as on older VSTM representations (Exp. 1). (2) When operating within capacity, subjects use the cue to prioritise where they initiate their memory search, rather than to discard uncued items (Exps. 2 and 3). (3) When capacity is exceeded, there is little benefit to top-down biasing relative to a neutral condition; however, unattended items are lost, with there being a substantial cost of invalid spatial cueing (Exp. 3). (4) These costs and benefits of orienting spatial attention differ across iconic memory and VSTM representations when VSTM capacity is exceeded (Exp. 4).

Spatial attention can bias search in visual short-term memory.

Whereas top-down attentional control is known to bias perceptual functions at many levels of stimulus analysis, its possible influence over memory-related functions remains uncharted. Our experiment combined behavioral measures and event-related potentials (ERPs) to test the ability of spatial orienting to bias functions associated with visual short-term memory (VSTM), and to shed light on the neural mechanisms involved. In particular, we investigated whether orienting attention to a spatial location within an array maintained in VSTM could facilitate the search for a specific remembered item. Participants viewed arrays of one, two or four differently colored items, followed by an informative spatial (100% valid) or uninformative neutral retro-cue (1500-2500 ms after the array), and later by a probe stimulus (500-1000 ms after the retro-cue). The task was to decide whether the probe stimulus had been present in the array. Behavioral results showed that spatial retro-cues improved both accuracy and response times for making decisions about the presence of the probe item in VSTM, and significantly attenuated performance decrements caused by increasing VSTM load. We also identified a novel ERP component (N3(RS)) specifically associated with searching for an item within VSTM. Paralleling the behavioral results, the amplitude and duration of the N3(RS) systematically increased with VSTM load in neutral retro-cue trials. When spatial retro-cues were provided, this "retro-search" component was absent. Our findings clearly show that the influence of top-down attentional biases extends to mnemonic functions, and, specifically, that searching for items within VSTM can be under flexible voluntary control.

Directing spatial attention in mental representations: Interactions between attentional orienting and working-memory load.

Orienting spatial attention to locations in the extrapersonal world has been intensively investigated during the past decades. Recently, it was demonstrated that it is also possible to shift attention to locations within mental representations held in working memory. This is an important issue, since the allocation of our attention is not only guided by external stimuli, but also by their internal representations and the expectations we build upon them. The present experiment used behavioural measures and event-related functional magnetic resonance imaging to investigate whether spatial orienting to mental representations can modulate the search and retrieval of information from working memory, and to identify the neural systems involved, respectively. Participants viewed an array of coloured crosses. Seconds after its disappearance, they were cued to locations in the array with valid or neutral cues. Subsequently, they decided whether a probe stimulus was presented in the array. The behavioural results indicated that orienting of spatial attention within working memory attenuates the well-known effect of decreasing performance when memory load is increased. So "internal" spatial orienting seems to highlight information or facilitate search within working memory, which leads to advantages in retrieval. Imaging enabled the separation of brain areas supporting spatial orienting functions from those sensitive to working-memory load. Orienting of spatial attention to the contents of working memory activated posterior parietal cortex bilaterally, the insula, and lateral and medial frontal cortices.

Orienting attention to locations in internal representations.

Three experiments investigated whether it is possible to orient selective spatial attention to internal representations held in working memory in a similar fashion to orienting to perceptual stimuli. In the first experiment, subjects were either cued to orient to a spatial location before a stimulus array was presented (pre-cue), cued to orient to a spatial location in working memory after the array was presented (retro-cue), or given no cueing information (neutral cue). The stimulus array consisted of four differently colored crosses, one in each quadrant. At the end of a trial, a colored cross (probe) was presented centrally, and subjects responded according to whether it had occurred in the array. There were equivalent patterns of behavioral costs and benefits of cueing for both pre-cues and retro-cues. A follow-up experiment used a peripheral probe stimulus requiring a decision about whether its color matched that of the item presented at the same location in the array. Replication of the behavioral costs and benefits of pre-cues and retro-cues in this experiment ruled out changes in response criteria as the only explanation for the effects. The third experiment used event-related potentials (ERPs) to compare the neural processes involved in orienting attention to a spatial location in an external versus an internal spatial representation. In this task, subjects responded according to whether a central probe stimulus occurred at the cued location in the array. There were both similarities and differences between ERPs to spatial cues toward a perception versus an internal spatial representation. Lateralized early posterior and later frontal negativities were observed for both pre- and retro-cues. Retro-cues also showed additional neural processes to be involved in orienting to an internal representation, including early effects over frontal electrodes.

Multiple mechanisms of selective attention: differential modulation of stimulus processing by attention to space or time.

Two studies compared the modulatory effects of orienting attention to spatial locations versus temporal intervals using event-related potentials (ERPs). In both experiments subjects performed attentional orienting tasks, which used identical stimuli in both spatial and temporal orienting conditions. The first experiment (N=16) used bilateral peripheral targets (7.5 degrees eccentricity) at two different time intervals (600, 1200 ms after cue onset). During spatial orienting a symbolic central cue predicted (75% probability) the spatial location (left, right) of the relevant target. No information was given about the probable target interval (short, long). In temporal orienting the cue predicted the target interval but not its location. Valid cueing produced significantly shorter reaction times in both the spatial and temporal orienting conditions. ERPs to identical, non-target stimulus arrays were analysed, to isolate endogenous attentional mechanisms. Spatial and temporal attention had distinct modulatory effects upon stimulus processing. Focused spatial attention affected the amplitude of early visual components. Modulation by temporal attention started later, and mainly affected potentials linked to decisions and responses. The second experiment (N=12) used unilateral target stimuli, and equated the probability of stimulus occurrence at short and long time intervals and at left or right of fixation. The results confirmed the distinct pattern of modulation of stimulus processing by spatial and temporal orienting. The optimisation of behaviour by attention can thus be achieved as a consequence of distinct modulatory processes, illustrating the flexibility of attentional functions in the human brain.

Involvement of the medial pallidum in focal myoclonic dystonia: A clinical and neurophysiological case study.

We successfully treated a patient with familial myoclonic dystonia (FMD), which primarily affected his neck muscles, with bilateral deep brain stimulation (DBS) to the medial pallidum, and investigated the role of the medial pallidum in FMD. A patient with FMD underwent bilateral implantation of DBS electrodes during which field potentials (FPs) in the medial pallidum and electromyograms (EMGs) from the affected neck muscles were recorded. The effects of high-frequency DBS to the medial pallidum on the FMD were also assessed by recording EMGs during and immediately after implantation, as well as 6 days and 8 weeks postoperatively. During spontaneous myoclonic episodes, increased FPs oscillating at 4 and 8 Hz were recorded from the medial pallidum; these correlated strongly with phasic EMG activity at the same frequencies in the contralateral affected muscles. The EMG activity was suppressed by stimulating the contralateral medial pallidum at 100 Hz during the operation and continuous bilateral DBS from an implanted stimulator abolished myoclonic activity even more effectively postoperatively. The phasic pallidal activity correlated with and led the myoclonic muscle activity, and the myoclonus was suppressed by bilateral pallidal DBS, suggesting that the medial pallidum was involved in the generation of the myoclonic activity. High-frequency DBS may suppress the myoclonus by desynchronising abnormal pallidal oscillations. This case study has significant clinical implications, because at present, there is no effective treatment for focal myoclonic dystonia.