Updating spatial working memory in a dynamic visual environment.

The present review describes recent developments regarding the role of the eye movement system in representing spatial information and keeping track of locations of relevant objects. First, we discuss the active vision perspective and why eye movements are considered crucial for perception and attention. The second part focuses on the question of how the oculomotor system is used to represent spatial attentional priority, and the role of the oculomotor system in maintenance of this spatial information. Lastly, we discuss recent findings demonstrating rapid updating of information across saccadic eye movements. We argue that the eye movement system plays a key role in maintaining and rapidly updating spatial information. Furthermore, we suggest that rapid updating emerges primarily to make sure actions are minimally affected by intervening eye movements, allowing us to efficiently interact with the world around us.

Rapid updating of spatial working memory across saccades.

Each time we make an eye movement, positions of objects on the retina change. In order to keep track of relevant objects their positions have to be updated. The situation becomes even more complex if the object is no longer present in the world and has to be held in memory. In the present study, we used saccadic curvature to investigate the time-course of updating a memorized location across saccades. Previous studies have shown that a memorized location competes with a saccade target for selection on the oculomotor map, which leads to saccades curving away from it. In our study participants performed a sequence of two saccades while keeping a location in memory. The trajectory of the second saccade was used to measure when the memorized location was updated after the first saccade. The results showed that the memorized location was rapidly updated with the eyes curving away from its spatial coordinates within 130 ms after the first eye movement. The time-course of updating was comparable to the updating of an exogenously attended location, and depended on how well the location was memorized.

Eye abduction reduces but does not eliminate competition in the oculomotor system.

Although it is well established that there is a tight coupling between covert attention and the eye movement system there is an ongoing controversy whether this relationship is functional. Previous studies demonstrated that disrupting the ability to execute an eye movement interferes with the allocation of covert attention. One technique that prevents the execution of an eye movement involves the abduction of the eye in the orbit while presenting the stimuli outside of the effective oculomotor range (Craighero, Nascimben, & Fadiga, 2004). Although eye abduction is supposed to disrupt activation of the oculomotor program responsible for the shift of covert attention, this crucial assumption has never been tested experimentally. In the present study we used saccadic curvature to examine whether eye abduction eliminates the target-distractor competition in the oculomotor system. We experimentally reduced the ability to execute saccades by abducting the eye by 30° (monocular vision). This way the peripheral part of the temporal hemifield was located outside the oculomotor range. Participants made a vertical eye movement while on some trials a distractor was shown either inside or outside of the oculomotor range. The curvature away from distractors located outside the oculomotor range was reduced, but not completely eliminated. This confirms that eye abduction influences the activation of the oculomotor program, but points to the fact that other forms of motor planning, such as head movements are also represented in the oculomotor system. The results are in line with the idea that covert attention is an emerging property of movement planning, but is not restricted to saccade planning.

The Role of the Oculomotor System in Updating Visual-Spatial Working Memory across Saccades.

Visual-spatial working memory (VSWM) helps us to maintain and manipulate visual information in the absence of sensory input. It has been proposed that VSWM is an emergent property of the oculomotor system. In the present study we investigated the role of the oculomotor system in updating of spatial working memory representations across saccades. Participants had to maintain a location in memory while making a saccade to a different location. During the saccade the target was displaced, which went unnoticed by the participants. After executing the saccade, participants had to indicate the memorized location. If memory updating fully relies on cancellation driven by extraretinal oculomotor signals, the displacement should have no effect on the perceived location of the memorized stimulus. However, if postsaccadic retinal information about the location of the saccade target is used, the perceived location will be shifted according to the target displacement. As it has been suggested that maintenance of accurate spatial representations across saccades is especially important for action control, we used different ways of reporting the location held in memory; a match-to-sample task, a mouse click or by making another saccade. The results showed a small systematic target displacement bias in all response modalities. Parametric manipulation of the distance between the to-be-memorized stimulus and saccade target revealed that target displacement bias increased over time and changed its spatial profile from being initially centered on locations around the saccade target to becoming spatially global. Taken together results suggest that we neither rely exclusively on extraretinal nor on retinal information in updating working memory representations across saccades. The relative contribution of retinal signals is not fixed but depends on both the time available to integrate these signals as well as the distance between the saccade target and the remembered location.

Can an early 24-hour EEG predict the response to the ketogenic diet? A prospective study in 34 children and adults with refractory epilepsy treated with the ketogenic diet.

PURPOSE: We examined whether early EEG changes in a 24-h EEG at 6 weeks of treatment were related to the later clinical response to the ketogenic diet (KD) in a 6-month period of treatment. METHODS: We examined 34 patients with heterogeneous epilepsy syndromes (21 children, 13 adults) and found 9 clinical responders (≥50% seizure reduction); this is a responder rate of 26%. We visually counted the interictal epileptic discharge index (IED index) in % during 2h of wakefulness and in the first hour of sleep (method 1), and also globally reviewed EEG changes (method 2), while blinded to the effect of the KD. RESULTS: At group level we saw a correlation between nocturnal reduction of IED-index at 6 weeks and seizure reduction in the follow-up period. A proportional reduction in IED index of 30% from baseline in the sleep EEG, was associated with being a responder to the diet (Pearson Chi-square p=0.04). EEG scoring method 2 observed a significantly larger proportion of patients with EEG-improvement in sleep in KD responders than in non-responders (p=0.03). At individual level, however, EEG changes did not correlate very strongly to the response to the diet, as IED reduction in sleep was also seen in 15% (method 1) to 26% (method 2) of the non-responders. CONCLUSION: Nocturnal reduction of IEDs is related to the response to the KD, however in daily clinical practice, an early EEG to predict seizure reduction should not be advised for individual patients.

Updating visual-spatial working memory during object movement.

Working memory enables temporary maintenance and manipulation of information for immediate access by cognitive processes. The present study investigates how spatial information stored in working memory is updated during object movement. Participants had to remember a particular location on an object which, after a retention interval, started to move. The question was whether the memorized location was updated with the movement of the object or whether after object movement it remained represented in retinotopic coordinates. We used saccade trajectories to examine how memorized locations were represented. The results showed that immediately after the object stopped moving, there was both a retinotopic and an object-centered representation. However, 200ms later, the activity at the retinotopic location decayed, making the memory representation fully object-centered. Our results suggest that memorized locations are updated from retinotopic to object-centered coordinates during, or shortly after object movement.