Martijn Meeter

Eye movement trajectories and what they tell us

In the last two decades, research has shown that eye movement trajectories can be modified by situational determinants. These modifications can inform us about the mechanisms that control eye movements and they can yield information about the oculomotor, memory and attention system that is not easily obtained via other sources. Eye movement trajectories can deviate either towards or away from elements in the visual field. We review the conditions in which these deviations are found and the mechanisms underlying trajectory deviations. It is argued that deviations towards an element are caused by the unresolved competition in the oculomotor system between elements in a visual scene. Deviations away from an element are mainly observed in situations in which top-down preparation can influence the target selection process, but the exact cause of such deviations remains unclear.

Consolidation of Long-Term Memory: Evidence and Alternatives

Memory loss in retrograde amnesia has long been held to be larger for recent periods than for remote periods, a pattern usually referred to as the Ribot gradient. One explanation for this gradient is consolidation of long-term memories. Several computational models of such a process have shown how consolidation can explain characteristics of amnesia, but they have not elucidated how consolidation must be envisaged. Here findings are reviewed that shed light on how consolidation may be implemented in the brain. Moreover, consolidation is contrasted with alternative theories of the Ribot gradient. Consolidation theory, multiple trace theory, and semantization can all handle some findings well but not others. Conclusive evidence for or against consolidation thus remains to be found.

Intertrial priming stemming from ambiguity: A new account of priming in visual search

Sequential effects are ubiquitous in experimental psychology. Within visual search, performance is often speeded when participants search for the same target twice in a row, as opposed to two different targets. Here, we investigate such intertrial priming. Two experiments show that factors influencing search processes affect the presence and size of intertrial priming: It is larger when there are fewer elements in the visual display, and larger when there is a salient distractor present than when the target is the only salient element in the display. A control experiment showed that these increased priming effects were not due to longer baseline RTs. These findings, it is argued, are inconsistent with theories that explain intertrial priming as resulting from either only faster visual selection, or from episodic retrieval of responses. Instead, we propose that ambiguity in the stimulus or task underlies the occurrence of intertrial priming.

Neural Mechanisms Underlying Probabilistic Category Learning in Normal Aging

Probabilistic category learning engages neural circuitry that includes the prefrontal cortex and caudate nucleus, two regions that show prominent changes with normal aging. However, the specific contributions of these brain regions are uncertain, and the effects of normal aging have not been examined previously in probabilistic category learning. In the present study, using a blood oxygenation level-dependent functional magnetic resonance imaging block design, 18 healthy young adults (mean age, 25.5 ± 2.6 years) and 15 older adults (mean age, 67.1 ± 5.3 years) were assessed on the probabilistic category learning “weather prediction” test. Whole-brain functional images acquired using a 1.5T scanner (General Electric, Milwaukee, WI) with gradient echo, echo planar imaging (3/1 mm; repetition time, 3000 ms; echo time, 50 ms) were analyzed using second-level random-effects procedures [SPM99 (Statistical Parametric Mapping)]. Young and older adults displayed equivalent probabilistic category learning curves, used similar strategies, and activated analogous neural networks, including the prefrontal and parietal cortices and the caudate nucleus. However, the extent of caudate and prefrontal activation was less and parietal activation was greater in older participants. The percentage correct and reaction time were mainly positively correlated with caudate and prefrontal activation in young individuals but positively correlated with prefrontal and parietal cortices in older individuals. Differential activation within a circumscribed neural network in the context of equivalent learning suggests that some brain regions, such as the parietal cortices, may provide a compensatory mechanism for healthy older adults in the context of deficient prefrontal cortex and caudate nuclei responses.

On the dissociation between compound and present/absent tasks in visual search: Intertrial priming is ambiguity driven

Visual search is speeded when the target-defining property (a feature- or dimension difference relative to the distractors) is repeated relative to when it changes. It is thought that automatic and implicit intertrial priming mechanisms underlie this effect. However, intertrial priming has been found to be less robust in compound search tasks (in which the response property is unrelated to the target-defining property) than in present/absent search tasks (in which the response is directly related to the presence of a target-defining property). This study explored the hypothesis that intertrial priming is dependent on the level of ambiguity in a task, with the present/absent task being inherently more ambiguous than the compound search task. The first three of five experiments further established the dissociation between the tasks and excluded alternative explanations. Intertrial priming was strong in present/absent and go/no-go tasks, but absent in compound and compound/absent tasks. The last two experiments supported the ambiguity hypothesis by introducing more uncertainty in the compound task, after which intertrial priming returned.

A competitive integration model of exogenous and endogenous eye movements

We present a model of the eye movement system in which the programming of an eye movement is the result of the competitive integration of information in the superior colliculi (SC). This brain area receives input from occipital cortex, the frontal eye fields, and the dorsolateral prefrontal cortex, on the basis of which it computes the location of the next saccadic target. Two critical assumptions in the model are that cortical inputs are not only excitatory, but can also inhibit saccades to specific locations, and that the SC continue to influence the trajectory of a saccade while it is being executed. With these assumptions, we account for many neurophysiological and behavioral findings from eye movement research. Interactions within the saccade map are shown to account for effects of distractors on saccadic reaction time (SRT) and saccade trajectory, including the global effect and oculomotor capture. In addition, the model accounts for express saccades, the gap effect, saccadic reaction times for antisaccades, and recorded responses from neurons in the SC and frontal eye fields in these tasks.

Effects of 5-HT on memory and the hippocampus: model and data.

5-Hydroxytryptamine (5-HT) transmission has been implicated in memory and in depression. Both 5-HT depletion and specific 5-HT agonists lower memory performance, while depression is also associated with memory deficits. The precise neuropharmacology and neural mechanisms underlying these effects are unknown. We used neural network simulations to elucidate the neuropharmacology and network mechanisms underlying 5-HT effects on memory. The model predicts that these effects are largely dependent on transmission over the 5-HT1A and 5-HT3 receptors, which regulate the selectivity of retrieval. It also predicts differential memory deficit profiles for 5-HT depletion and overactivation. The latter predictions were confirmed in studies with healthy and depressed participants undergoing acute tryptophan depletion or ipsipirone challenge. The results suggest that the memory impairments in depressed subjects may be related to 5-HT undertransmission, and support the notion that 5-HT1A agonists ameliorate memory deficits in depression.

The limits of top-down control of visual attention

The extent to which spatial selection is driven by the goals of the observer and by the properties of the environment is one of the major issues in the field of visual attention. Here we review recent experimental evidence from behavioral and eye movement studies suggesting that top-down control has temporal and spatial limits. More specifically, we argue that the first feedforward sweep of information is bottom-up, and that top-down control can influence selection only after the sweep is completed. In addition, top-down control can limit spatial selection through adjusting the size of attentional window, an area of visual space which receives priority in information sampling. Finally, we discuss the evidence found using brain imaging techniques for top-down control in an attempt to reconcile it with behavioral findings. We conclude by discussing theoretical implications of these results for the current models of visual selection.

Neural Correlates of Probabilistic Category Learning in Patients with Schizophrenia

Functional neuroimaging studies of probabilistic category learning in healthy adults report activation of cortical-striatal circuitry. Based on previous findings of normal learning rate concurrent with an overall performance deficit in patients with schizophrenia, we hypothesized that relative to healthy adults, patients with schizophrenia would display preserved caudate nucleus and abnormal prefrontal cortex activation during probabilistic category learning. Forty patients with schizophrenia receiving antipsychotic medication and 25 healthy participants were assessed on interleaved blocks of probabilistic category learning and control tasks while undergoing blood oxygenation level-dependent functional magnetic resonance imaging. In addition to the whole sample of patients with schizophrenia and healthy adults, a subset of patients and healthy adults matched for good learning was also compared. In the whole sample analysis, patients with schizophrenia displayed impaired performance in conjunction with normal learning rate relative to healthy adults. The matched comparison of patients and healthy adults classified as good learners revealed greater caudate and dorsolateral prefrontal cortex activity in the healthy adults and greater activation in a more rostral region of the dorsolateral prefrontal, cingulate, parahippocampal and parietal cortex in patients. These results demonstrate that successful probabilistic category learning can occur in the absence of normal frontal-striatal function. Based on analyses of the patients and healthy adults matched on learning and performance, a minority of patients with schizophrenia achieve successful probabilistic category learning and performance levels through differential activation of a circumscribed neural network which suggests a compensatory mechanism in patients showing successful learning.

Time-dependent effects of cortisol on the contextualization of emotional memories

BACKGROUND The inability to store fearful memories into their original encoding context is considered to be an important vulnerability factor for the development of anxiety disorders like posttraumatic stress disorder. Altered memory contextualization most likely involves effects of the stress hormone cortisol, acting via receptors located in the memory neurocircuitry. Cortisol via these receptors induces rapid nongenomic effects followed by slower genomic effects, which are thought to modulate cognitive function in opposite, complementary ways. Here, we targeted these time-dependent effects of cortisol during memory encoding and tested subsequent contextualization of emotional and neutral memories. METHODS In a double-blind, placebo-controlled design, 64 men were randomly assigned to one of three groups: 1) received 10 mg hydrocortisone 30 minutes (rapid cortisol effects) before a memory encoding task; 2) received 10 mg hydrocortisone 210 minutes (slow cortisol) before a memory encoding task; or 3) received placebo at both times. During encoding, participants were presented with neutral and emotional words in unique background pictures. Approximately 24 hours later, context dependency of their memories was assessed. RESULTS Recognition data revealed that cortisols rapid effects impair emotional memory contextualization, while cortisols slow effects enhance it. Neutral memory contextualization remained unaltered by cortisol, irrespective of the timing of the drug. CONCLUSIONS This study shows distinct time-dependent effects of cortisol on the contextualization of specifically emotional memories. The results suggest that rapid effects of cortisol may lead to impaired emotional memory contextualization, while slow effects of cortisol may confer protection against emotional memory generalization.