Dynamic neuroplasticity after human prefrontal cortex damage.

Memory and attention deficits are common after prefrontal cortex (PFC) damage, yet people generally recover some function over time. Recovery is thought to be dependent upon undamaged brain regions, but the temporal dynamics underlying cognitive recovery are poorly understood. Here, we provide evidence that the intact PFC compensates for damage in the lesioned PFC on a trial-by-trial basis dependent on cognitive load. The extent of this rapid functional compensation is indexed by transient increases in electrophysiological measures of attention and memory in the intact PFC, detectable within a second after stimulus presentation and only when the lesioned hemisphere is challenged. These observations provide evidence supporting a dynamic and flexible model of compensatory neural plasticity.

Recognition memory of newly learned faces.

We used event-related fMRI to study recognition memory of newly learned faces. Caucasian subjects memorized unfamiliar, neutral and happy South Korean faces and 4 days later performed a memory retrieval task in the MR scanner. We predicted that previously seen faces would be recognized faster and more accurately and would elicit stronger neural activation than novel faces. Consistent with our hypothesis, novel faces were recognized more slowly and less accurately than previously seen faces. We found activation in a distributed cortical network that included face-responsive regions in the visual cortex, parietal and prefrontal regions, and the hippocampus. Within all regions, correctly recognized, previously seen faces evoked stronger activation than novel faces. Additionally, in parietal and prefrontal cortices, stronger activation was observed during correct than incorrect trials. Finally, in the hippocampus, false alarms to happy faces elicited stronger responses than false alarms to neutral faces. Our findings suggest that face recognition memory is mediated by stimulus-specific representations stored in extrastriate regions; parietal and prefrontal regions where old and new items are classified; and the hippocampus where veridical memory traces are recovered.

Recognition memory is modulated by visual similarity.

We used event-related fMRI to test whether recognition memory depends on visual similarity between familiar prototypes and novel exemplars. Subjects memorized portraits, landscapes, and abstract compositions by six painters with a unique style, and later performed a memory recognition task. The prototypes were presented with new exemplars that were either visually similar or dissimilar. Behaviorally, novel, dissimilar items were detected faster and more accurately. We found activation in a distributed cortical network that included face- and object-selective regions in the visual cortex, where familiar prototypes evoked stronger responses than new exemplars; attention-related regions in parietal cortex, where responses elicited by new exemplars were reduced with decreased similarity to the prototypes; and the hippocampus and memory-related regions in parietal and prefrontal cortices, where stronger responses were evoked by the dissimilar exemplars. Our findings suggest that recognition memory is mediated by classification of novel exemplars as a match or a mismatch, based on their visual similarity to familiar prototypes.

Temporal kinetics of prefrontal modulation of the extrastriate cortex during visual attention.

Single-unit, event-related potential (ERP), and neuroimaging studies have implicated the prefrontal cortex (PFC) in top-down control of attention and working memory. We conducted an experiment in patients with unilateral PFC damage (n = 8) to assess the temporal kinetics of PFC-extrastriate interactions during visual attention. Subjects alternated attention between the left and the right hemifields in successive runs while they detected target stimuli embedded in streams of repetitive task-irrelevant stimuli (standards). The design enabled us to examine tonic (spatial selection) and phasic (feature selection) PFC-extrastriate interactions. PFC damage impaired performance in the visual field contralateral to lesions, as manifested by both larger reaction times and error rates. Assessment of the extrastriate P1 ERP revealed that the PFC exerts a tonic (spatial selection) excitatory input to the ipsilateral extrastriate cortex as early as 100 msec post stimulus delivery. The PFC exerts a second phasic (feature selection) excitatory extrastriate modulation from 180 to 300 msec, as evidenced by reductions in selection negativity after damage. Finally, reductions of the N2 ERP to target stimuli supports the notion that the PFC exerts a third phasic (target selection) signal necessary for successful template matching during postselection analysis of target features. The results provide electrophysiological evidence of three distinct tonic and phasic PFC inputs to the extrastriate cortex in the initial few hundred milliseconds of stimulus processing. Damage to this network appears to underlie the pervasive deficits in attention observed in patients with prefrontal lesions.

Attention capture by auditory significant stimuli: semantic analysis follows attention switching.

Event-related potentials (ERPs) were recorded from the scalp to investigate a long-standing controversy in auditory attention research, namely when the 'breakthrough of the unattended' takes place in the human brain. Nine subjects classified visual stimuli appearing 300 ms after task-irrelevant standard tones (80%, i.e. P = 0.8) or novel sounds (20%, i.e. P = 0.2) into odd/even categories. After the recording session, subjects scored the novel sounds as to whether they had any particular meaning (identifiable) or were perceived as a burst of noise (non-identifiable), and performance and ERPs were analysed according to this classification. A control condition, in which the visual stimuli were presented with no sounds, showed that subjects covertly monitored the task-irrelevant sounds during visual task-performance, and a further condition, in which the auditory and visual stimuli appeared regardless of each other, made it possible to trace the processing of the distracters during allocation of attention outside the auditory environment. Results yielded identical N1-enhancement for the two types of novel sounds, indicating similar attention switching triggered to these two types of unexpected sounds. However, there was a stronger orientating of attention towards identifiable novel sounds, as indicated both by behavioural distraction and by larger novelty-P3. Furthermore, this stronger orientating of attention was due to the sounds being contingent on the visual stimuli, as no increase in novelty-P3 to identifiable novel sounds was observed in the control condition, in which the sounds occurred outside the attentional set. Therefore, provided that the N1-enhancement reflects a call for focal attention, and novelty-P3 the effective orientating of attention towards the eliciting sounds, the present results suggest that semantic analysis of significant sounds occurs after a transitory switch of attention towards the eliciting stimuli. Moreover, as the novelty-P3 increase in amplitude was observed only when subjects covertly monitored the sounds, the present data suggest that semantic analysis of irrelevant sounds depends on the top-down cognitive influences of the attentional set.

Spatiotemporal dynamics of the auditory novelty-P3 event-related brain potential.

The spatiotemporal dynamics of the cerebral network involved in novelty processing was studied by means of scalp current density (SCD) analysis of the novelty P3 (nP3) event-related brain potential (ERP). ERPs were recorded from 30 scalp electrodes at the occurrence of novel unpredictable environmental sounds during the performance of a visual discrimination task. Increased SCD was observed at left frontotemporal (FT3), bilateral temporoparietal (TP3 and TP4) and prefrontal locations (F8-F4 and F7-F3), suggesting novelty-P3 generators located in the left auditory cortex, and bilaterally in temporoparietal and prefrontal association regions. Additional increased SCD was found at a central location (Cz) and at superior parietal locations (P3-Pz-P4). The SCD of the nP3 was therefore generated at three successive, partially overlapping, stages of neuroelectric activation. At the central location, SCD started to be significant before the onset of the nP3 waveform, contributing solely to its early phase. At temporoparietal and left frontotemporal locations, nP3 electrophysiological activity was characterized by sustained current density, starting at about 210 ms and continuing during the full latency range of the response, including its early and late phases. At its late phase, the nP3 was characterized by sharp phasic current density at prefrontal and superior parietal locations, starting at about 290 ms and vanishing at around 385 ms. Taken together, these results provide the first evidence of the cerebral spatio-temporal dynamics underlying novelty processing.

Electrophysiological evidence of abnormal activation of the cerebral network of involuntary attention in alcoholism.

OBJECTIVE: Increased distractibility is a common impairment in alcoholism, but objective evidence has remained elusive. Here, a task designed to investigate with event-related brain potentials (ERPs) the neural mechanism underlying distraction was used to show abnormal involuntary orienting of attention in chronic alcoholism. METHODS: Fifteen alcoholics and 17 matched healthy controls were instructed to ignore auditory stimuli while concentrating in the discrimination of immediately following visual stimuli. The auditory sequences contained repetitive standard tones occasionally replaced by deviant tones of slightly higher frequency, or by complex novel sounds. RESULTS: Deviant tones and novel sounds distracted visual performance, i.e. increased reaction time to visual stimuli, similarly in patients and controls. Compared to controls, however, alcoholics showed ERP abnormalities, i.e. enhanced P3a amplitudes over the left frontal region, and a positive posterior deflection instead of the frontally distributed reorienting negativity (RON). CONCLUSIONS: The enhanced P3a to novelty and subsequent positive wave instead of RON in alcoholics suggests encoding into working memory of task-irrelevant auditory events and provides neurophysiological markers of impaired involuntary attention mechanisms in chronic alcoholism.

An electrophysiological and behavioral investigation of involuntary attention towards auditory frequency, duration and intensity changes.

We measured behavior and event-related brain potentials (ERPs) in 12 subjects performing on an audio-visual distraction paradigm to investigate the cerebral mechanisms of involuntary attention towards stimulus changes in the acoustic environment. Subjects classified odd/even numbers presented on a computer screen 300 ms after the occurrence of a task-irrelevant auditory stimulus, by pressing the corresponding response button. Auditory stimuli were standard tones (600 Hz, 200 ms, 85 dB; P=0.8) or deviant tones (P=0.2), these differing from the standard either in frequency (700 Hz), duration (50 ms) or intensity (79 dB), in separate blocks. In comparison to performance to visual stimuli following the standard tones, reaction time increased by 24 ms (F(1,11)=10.91, P<0.01) and hit rate decreased by 4.6% (F(1,11)=35.47, P<0.001) to visual stimuli following the deviant tones, indicating behavioral distraction. ERPs revealed the mismatch negativity (MMN) elicited to deviant tones, which was larger for the duration deviant than for the frequency and intensity deviants (F(2,22)=19.43, P<0.001, epsilon =0.83), and which had different scalp distribution for all three deviant conditions (F(16,176)=2.40, P<0.05, epsilon =0.12). As the shorter duration and softer intensity deviant tones were unlikely to engage fresh neurons responding to their specific physical features, the present results indicate that a genuine change detection mechanism is involved in triggering attention switching towards sound changes, and suggest a largely distributed neural network of the auditory cortex underlying such involuntary attention switching.