Fluid reasoning, or the capacity to think logically and solve novel problems, is central to the development of human cognition, but little is known about the underlying neural changes. During the acquisition of event-related fMRI data, children aged 6-13 (N = 16) and young adults (N = 17) performed a task in which they were asked to identify semantic relationships between drawings of common objects. On semantic problems, participants indicated which of five objects was most closely semantically related to a cued object. On analogy problems, participants solved a visual propositional analogy ( e. g., shoe is to foot as glove is to.?) by indicating which of four objects would complete the problem; these problems required integration of two semantic relations, or relational integration. Our prior research on analogical reasoning in adults implicated left anterior ventrolateral prefrontal cortex (VLPFC) in the controlled retrieval of individual semantic relationships, and rostrolateral prefrontal cortex (RLPFC) in relational integration. In this study, age-related changes in the recruitment of VLPFC, temporal cortex, and other cortical regions were observed during the retrieval of individual semantic relations. In contrast, age-related changes in RLPFC function were observed during relational integration. Children aged 6-13 engage RLPFC too late in the analogy trials to influence their behavioral responses, suggesting that important changes in RLPFC function take place during adolescence.
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 (N3RS) specifically associated with searching for an item within VSTM. Paralleling the behavioral results, the amplitude and duration of the N3RS 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.
The Nucleus accumbens (Nacc) has been proposed to act as a limbic-motor interface. Here, using invasive intraoperative recordings in an awake patient suffering from obsessive-compulsive disease (OCD), we demonstrate that its activity is modulated by the quality of performance of the subject in a choice reaction time task designed to tap action monitoring processes. Action monitoring, that is, error detection and correction, is thought to be supported by a system involving the dopaminergic midbrain, the basal ganglia, and the medial prefrontal cortex. In surface electrophysiological recordings, action monitoring is indexed by an error-related negativity ( ERN) appearing time-locked to the erroneous responses and emanating from the medial frontal cortex. In preoperative scalp recordings the patient's ERN was found to be significantly increased compared to a large (n = 83) normal sample, suggesting enhanced action monitoring processes. Intraoperatively, error-related modulations were obtained from the Nacc but not from a site 5 mm above. Importantly, cross-correlation analysis showed that error-related activity in the Nacc preceded surface activity by 40 ms. We propose that the Nacc is involved in action monitoring, possibly by using error signals from the dopaminergic midbrain to adjust the relative impact of limbic and prefrontal inputs on frontal control systems in order to optimize goal-directed behavior.
This study aimed to clarify the neural substrates of behavioral switch and restart costs in intermittently instructed task-switching paradigms. Event-related potentials (ERPs) were recorded while participants were intermittently cued to switch or repeat their categorization rule (Switch task), or else they performed two perceptually identical control conditions (NoGo and Oddball). The three tasks involved different task-sets with distinct stimulus-response associations in each, but identical visual stimulation, consisting of frequent colored shapes (p = 0.9) and randomly interspersed infrequent black shapes (p = 0.1; '+' and 'X' symbols). Behavioral restart costs were observed in the first target responses following all black shapes in the Switch and NoGo tasks - but not in the Oddball task - and corresponded with enhanced fronto-centrally distributed early cue-locked P3 activity (peak latency 325-375 ms post-cue onset at the vertex). In turn, behavioral switch costs were associated with larger late cue-locked P3 amplitudes in the Switch task only (peak latency 400 - 450 ms post-cue onset at mid-parietal sites). Together with our information theoretical estimations, ERP results suggested that restart and switch costs indexed two neural mechanisms related to the preparatory resolution of uncertainty: ( 1) the intermittent re-activation of task-set information, and (2) the updating of stimulus-response mappings within an active task set, as indexed by early and late cue-locked P3 activations, respectively. In contrast, target-locked P3 activations reflected a functionally distinct mechanism related to the implementation of task-set information. We conclude that task-switching costs consist of both switch-specific and switch-unspecific processes during the preparation and execution stages of task performance.
In this article we review research in our laboratory on auditory hallucinations using behavioral and MRI measure. The review consists of both previously published and new data that for the first time is presented together in a cohesive way. Auditory hallucinations are among the most common symptoms in schizophrenia, affecting more than 70% of the patients. We here advance the hypothesis that auditory hallucinations are internally generated speech perceptions that are lateralized to the left temporal lobe, in the peri-Sylvian region. From this we predict that hallucinating patients should have problems identifying a simultaneously presented external speech sound, as measured through performance on the dichotic listening (DL) paradigm with consonant-vowel syllables, since this technique lateralizes the stimulus input. Across a series of behavioral experiments, we have shown that patients with schizophrenia who experience frequent auditory hallucinations fail to demonstrate an expected right ear advantage on the dichotic listening test. Absence of a right ear advantage is indicative of a functional deficit in the left peri-Sylvian region. The results also revealed that patients with ongoing auditory hallucinations were more impaired than patients with previous hallucinations, and that a higher score on the hallucination item in a standard symptom rating scale (BPRS) correlated negatively with number of correct reports for the right ear stimulus. Moreover, we have found that schizophrenia patients fail to shift attention to the left ear stimulus, when explicitly instructed to focus on the right or left ear stimulus only, thus showing a deficit in inhibition of attention and response-inhibition. The behavioral DL data are substantiated in two MR morphometry studies that revealed significant reductions in grey matter density in the left peri-Sylvian region in hallucinating patients, and patients with reduced left temporal lobe grey matter density. Hallucinating patients also failed to show a right ear advantage in the dichotic listening test. Ongoing fMRI studies are focused on the underlying synaptic and molecular mechanisms by investigating the effects of the glutamate antagonist drug memantine on auditory perception and speech lateralization, and examination of temporal cortex-specific gene expression in the left peri-Sylvian region.
This study examined the performance of 41 patients with focal prefrontal cortical lesions and 38 healthy controls on a task-switching procedure. Three different conditions were evaluated: single tasks without switches and two switching tasks with the currently relevant task signalled either 1500 ms ( Long Cue) or 200 ms ( Short Cue) before the stimulus. Patients with Superior Medial lesions showed both a general slowing of reaction time (RT) and a significantly increased switch cost as measured by RT. No other prefrontal group showed this increased reaction time switch cost. Increased error rates in the switching conditions, on the other hand, were observed in patients with Inferior Medial lesions and, to a lesser extent, ones with Superior Medial lesions. Patients with left dorsolateral lesions (9/46v) showed slower learning of the task as indicated by a high error rate early on. Several different processes are involved in task-switching and these are selectively disrupted by lesions to specific areas of the frontal lobes.
The world is an unpredictable place, presenting challenges that fluctuate from moment to moment. However, the neural systems for responding to such challenges are far from fully understood. Using fMRI, we studied an audiovisual task in which the trials' difficulty and onset times varied unpredictably. Two regions were found to increase their activation for challenging trials, with their activities strongly correlated: right frontal cortex and the brainstem. The frontal area matched regions found in previous human studies of cognitive control, and activated in a graded manner with increasing task difficulty. The brainstem responded only to the most difficult trials, showing a phasic activity pattern paralleling locus coeruleus recordings in monkeys. These results reveal a bridge between animal and human studies, and suggest interacting roles for the brainstem and right frontal cortex: the brainstem may signal that an attentional challenge is occurring, while right frontal cortex allocates cognitive resources in response.
We examine the relative timing of numerous brain regions involved in human decisions that are based on external criteria, learned information, personal preferences, or unconstrained internal considerations. Using magnetoencephalography (MEG) and advanced signal analysis techniques, we were able to non-invasively reconstruct oscillations of distributed neural networks in the high-gamma frequency band (60-150 Hz). The time course of the observed neural activity suggested that two-alternative forced choice tasks are processed in four overlapping stages: processing of sensory input, option evaluation, intention formation, and action execution. Visual areas are activated first, and show recurring activations throughout the entire decision process. The temporo-occipital junction and the intraparietal sulcus are active during evaluation of external values of the options, 250-500 ms after stimulus presentation. Simultaneously, personal preference is mediated by cortical midline structures. Subsequently, the posterior parietal and superior occipital cortices appear to encode intention, with different subregions being responsible for different types of choice. The cerebellum and inferior parietal cortex are recruited for internal generation of decisions and actions, when all options have the same value. Action execution was accompanied by activation peaks in the contralateral motor cortex. These results suggest that high-gamma oscillations as recorded by MEG allow a reliable reconstruction of decision processes with excellent spatiotemporal resolution.
In everyday life, consciously or not, we are constantly disentangling the multiple auditory sources contributing to our acoustical environment. To better understand the neural mechanisms involved in concurrent sound processing, we manipulated sound onset asynchrony to induce the segregation or grouping of two concurrent sounds. Each sound consisted of amplitude-modulated tones at different carrier and modulation frequencies, allowing a cortical tagging of each sound. Electrophysiological recordings were carried out in epileptic patients with pharmacologically resistant partial epilepsy, implanted with depth electrodes in the temporal cortex. Patients were presented with the stimuli while they performed an auditory distracting task. We found that transient and steady-state evoked responses, and induced gamma oscillatory activities were enhanced in the case of onset synchrony. These effects were mainly located in the Heschl's gyrus for steady-state responses whereas they were found in the lateral superior temporal gyrus for evoked transient responses and induced gamma oscillations. They can be related to distinct neural mechanisms such as frequency selectivity and habituation. These results in the auditory cortex provide an anatomically refined description of the neurophysiological components which might be involved in the perception of concurrent sounds.
Human perception of faces is widely believed to rely on automatic processing by a domain-specific, modular component of the visual system. Scalp-recorded event-related potential (ERP) recordings indicate that faces receive special stimulus processing at around 170 ms poststimulus onset, in that faces evoke an enhanced occipital negative wave, known as the N170, relative to the activity elicited by other visual objects. As predicted by modular accounts of face processing, this early face-specific N170 enhancement has been reported to be largely immune to the influence of endogenous processes such as task strategy or attention. However, most studies examining the influence of attention on face processing have focused on non-spatial attention, such as object-based attention, which tend to have longer-latency effects. In contrast, numerous studies have demonstrated that visual spatial attention can modulate the processing of visual stimuli as early as 80 ms poststimulus-substantially earlier than the N170. These temporal characteristics raise the question of whether this initial face-specific processing is immune to the influence of spatial attention. This question was addressed in a dual-visual-stream ERP study in which the influence of spatial attention on the face-specific N170 could be directly examined. As expected, early visual sensory responses to all stimuli presented in an attended location were larger than responses evoked by those same stimuli when presented in an unattended location. More importantly, a significant face-specific N170 effect was elicited by faces that appeared in an attended location, but not in an unattended one. In summary, early face-specific processing is not automatic, but rather, like other objects, strongly depends on endogenous factors such as the allocation of spatial attention. Moreover, these findings underscore the extensive influence that top-down attention exercises over the processing of visual stimuli, including those of high natural salience.
We used magneto-encephalography (MEG) to measure visually evoked activity in healthy volunteers performing saccadic eye movements to visual targets. The neuromagnetic activity was analyzed from regions of cortical activation identified in separate functional magnetic resonance imaging (fMRI) studies. The latency of visual responses significantly increased from the Middle Temporal region (MT+) to the Intraparietal Sulcus (IPS) to the Frontal Eye Field (FEF), and their amplitude was greater in the hemisphere contralateral to the visual target. Trial-to-trial variability of oculomotor reaction times correlated with visual response latency across cortical areas. These results support a feedforward recruitment of oculomotor cortical centers by visual information, and a model in which behavioral variability depends on variability at different neural stages of processing.
Cues that direct selective attention to a spatial location have been observed to increase baseline neural activity in visual areas that represent a to-be-attended stimulus location. Analogous attention-related baseline shifts have also been observed in response to attention-directing cues for non-spatial stimulus features. It has been proposed that baseline shifts with preparatory attention may serve as the mechanism by which attention modulates the responses to subsequent visual targets that match the attended location or feature. Using functional MRI, we localized color- and motion-sensitive visual areas in individual subjects and investigated the relationship between cue-induced baseline shifts and the subsequent attentional modulation of task-relevant target stimuli. Although attention-directing cues often led to increased background neural activity in feature specific visual areas, these increases were not correlated with either behavior in the task or subsequent attentional modulation of the visual targets. These findings cast doubt on the hypothesis that attention-related shifts in baseline neural activity result in selective sensory processing of visual targets during feature-based selective attention.
Although positron emission tomography ( PET) and the aromatic L-amino acid decarboxylase (AADC) tracer 6-[F-18]fluoro-L-m-tyrosine (FMT) has been used to assess the integrity of the presynaptic dopamine system in the brain, relatively little has been published in terms of brain FMT uptake values especially for normal human subjects. Twelve normal volunteer subjects were scanned using PET and FMT to determine the range of normal striatal uptake values using Patlak graphical analysis. For comparison, seven adult rhesus monkeys were studied and the data analyzed in the same way. A subset of monkeys that were treated with a unilateral intracarotid artery infusion of the dopamine neurotoxin MPTP showed an 87% decrease in striatal FMT uptake. These findings support the use of PET and FMT to image AADC distribution in both normal and diseased brains using Patlak graphical analysis and tissue input functions.
The formation of coherent percepts requires grouping together spatio-temporally disparate sensory inputs. Two major questions arise: (1) is awareness necessary for this process; and (2) can non-conscious elements of the sensory input be grouped into a conscious percept? To address this question, we tested two patients suffering from severe left auditory extinction following right hemisphere damage. In extinction, patients are unaware of the presence of left side stimuli when they are presented simultaneously with right side stimuli. We used the 'scale illusion' to test whether extinguished tones on the left can be incorporated into the content of conscious awareness. In the scale illusion, healthy listeners obtain the illusion of distinct melodies, which are the result of grouping of information from both ears into illusory auditory streams. We show that the two patients were susceptible to the scale illusion while being consciously unaware of the stimuli presented on their left. This suggests that awareness is not necessary for auditory grouping and non-conscious elements can be incorporated into a conscious percept.