Memory encoding for important information can be enhanced both by reward anticipation and by intentional strategies. These effects are hypothesized to depend on distinct neural mechanisms, yet prior work has provided only limited evidence for their separability. We aimed to determine whether reward-driven and strategic mechanisms for prioritizing important information are separable, even if they may also interact. We examined the joint operation of both mechanisms using fMRI measures of brain activity. Participants learned abstract visual images in a value-directed recognition paradigm. On each trial, two novel images were presented simultaneously in different screen quadrants, one arbitrarily designated as high point value and one as low value. Immediately after each block of 16 study trials, the corresponding point rewards could be obtained in a test of item recognition and spatial location memory. During encoding trials leading to successful subsequent memory, especially of high-value images, increased activity was observed in dorsal frontoparietal and lateral occipitotemporal cortex. Furthermore, activity in a network associated with reward was higher during encoding when any image, of high or low value, was subsequently remembered. Functional connectivity between right medial temporal lobe and right ventral tegmental area, measured via psychophysiological interaction, was also greater during successful encoding regardless of value. Strategic control of memory, as indexed by successful prioritization of the high-value image, affected activity in dorsal posterior parietal cortex as well as connectivity between this area and right lateral temporal cortex. These results demonstrate that memory can be strengthened by separate neurocognitive mechanisms for strategic control versus reward-based enhancement of processing.

Language input is highly variable; phonological, lexical, and syntactic features vary systematically across different speakers, geographic regions, and social contexts. Previous evidence shows that language users are sensitive to these contextual changes and that they can rapidly adapt to local regularities. For example, listeners quickly adjust to accented speech, facilitating comprehension. It has been proposed that this type of adaptation is a form of implicit learning. This study examined a similar type of adaptation, syntactic adaptation, to address two issues: (1) whether language comprehenders are sensitive to a subtle probabilistic contingency between an extraneous feature (font color) and syntactic structure and (2) whether this sensitivity should be attributed to implicit learning. Participants read a large set of sentences, 40% of which were garden-path sentences containing temporary syntactic ambiguities. Critically, but unbeknownst to participants, font color probabilistically predicted the presence of a garden-path structure, with 75% of garden-path sentences (and 25% of normative sentences) appearing in a given font color. ERPs were recorded during sentence processing. Almost all participants indicated no conscious awareness of the relationship between font color and sentence structure. Nonetheless, after sufficient time to learn this relationship, ERPs time-locked to the point of syntactic ambiguity resolution in garden-path sentences differed significantly as a function of font color. End-of-sentence grammaticality judgments were also influenced by font color, suggesting that a match between font color and sentence structure increased processing fluency. Overall, these findings indicate that participants can implicitly detect subtle co-occurrences between physical features of sentences and abstract, syntactic properties, supporting the notion that implicit learning mechanisms are generally operative during online language processing.

Frequency-following and frequency-doubling neurons are ubiquitous in both striate and extrastriate visual areas. However, responses from these two types of neural populations have not been effectively compared in humans because previous EEG studies have not successfully dissociated responses from these populations. We devised a light–dark flicker stimulus that unambiguously distinguished these responses as reflected in the first and second harmonics in the steady-state visual evoked potentials. These harmonics revealed the spatial and functional segregation of frequency-following (the first harmonic) and frequency-doubling (the second harmonic) neural populations. Spatially, the first and second harmonics in steady-state visual evoked potentials exhibited divergent posterior scalp topographies for a broad range of EEG frequencies. The scalp maximum was medial for the first harmonic and contralateral for the second harmonic, a divergence not attributable to absolute response frequency. Functionally, voluntary visual–spatial attention strongly modulated the second harmonic but had negligible effects on the simultaneously elicited first harmonic. These dissociations suggest an intriguing possibility that frequency-following and frequency-doubling neural populations may contribute complementary functions to resolve the conflicting demands of attentional enhancement and signal fidelity—the frequency-doubling population may mediate substantial top–down signal modulation for attentional selection, whereas the frequency-following population may simultaneously preserve relatively undistorted sensory qualities regardless of the observer's cognitive state.

Familiarity and recollection are qualitatively different explicit-memory phenomena evident during recognition testing. Investigations of the neurocognitive substrates of familiarity and recollection, however, have typically disregarded implicit-memory processes likely to be engaged during recognition tests. We reasoned that differential neural responses to old and new items in a recognition test may reflect either explicit or implicit memory. Putative neural correlates of familiarity in prior experiments, for example, may actually reflect contamination by implicit memory. In two experiments, we used obscure words that subjects could not formally define to tease apart electrophysiological correlates of familiarity and one form of implicit memory, conceptual priming. In Experiment 1, conceptual priming was observed for words only if they elicited meaningful associations. In Experiment 2, two distinct neural signals were observed in conjunction with familiarity-based recognition: late posterior potentials for words that both did and did not elicit meaningful associations and FN400 potentials only for the former. Given that symbolic meaning is a prerequisite for conceptual priming, the combined results specifically link late posterior potentials and FN400 potentials with familiarity and conceptual priming, respectively. These findings contradict previous interpretations of FN400 potentials as generic signals of familiarity and show that repeated stimuli in recognition tests can engender facilitated processing of conceptual information in addition to retrieval processing that leads to the awareness of memory retrieval. The different characteristics of the electrical markers of these two types of process further underscore the biological validity of the distinction between implicit memory and explicit memory.

Stenberg et al. argued that FN400 brain potentials index familiarity rather than conceptual priming. Their data from a test of name recognition showed that both familiarity and FN400s were influenced by frequency but not fame, whereas separate behavioral measures of priming were influenced by fame but not frequency. However, this apparent dissociation was gravely weakened by confounds in task demands and inadequate behavioral measures of priming. Although Stenberg et al. failed to provide evidence suitable for disentangling neural correlates of familiarity from those of conceptual priming, an analysis of their report can be used to highlight difficulties that remain to be surmounted to understand recognition and the neural events that signal distinct memory functions engaged during recognition.

Affective judgments can often be influenced by emotional information people unconsciously perceive, but the neural mechanisms responsible for these effects and how they are modulated by individual differences in sensitivity to threat are unclear. Here we studied subliminal affective priming by recording brain potentials to surprise faces preceded by 30-msec happy or fearful prime faces. Participants showed valence-consistent changes in affective ratings of surprise faces, although they reported no knowledge of prime-face expressions, nor could they discriminate between prime-face expressions in a forced-choice test. In conjunction with the priming effect on affective evaluation, larger occipital P1 potentials at 145–175 msec were found with fearful than with happy primes, and source analyses implicated the bilateral extrastriate cortex in this effect. Later brain potentials at 300–400 msec were enhanced with happy versus fearful primes, which may reflect differential attentional orienting. Personality testing for sensitivity to threat, especially social threat, was also used to evaluate individual differences potentially relevant to subliminal affective priming. Indeed, participants with high trait anxiety demonstrated stronger affective priming and greater P1 differences than did those with low trait anxiety, and these effects were driven by fearful primes. Results thus suggest that unconsciously perceived affective information influences social judgments by altering very early perceptual analyses, and that this influence is accentuated to the extent that people are oversensitive to threat. In this way, perception may be subject to a variety of influences that govern social preferences in the absence of concomitant awareness of such influences.

Studies in healthy individuals and split-brain patients have shown that the representation of facial information from the left visual field (LVF) is better than the representation of facial information from the right visual field (RVF). To investigate the neurophysiological basis of this LVF superiority in face perception, we recorded event-related potentials (ERPs) to centrally presented face stimuli in which relevant facial information is present bilaterally (B faces) or only in the left (L faces) or the right (R faces) visual field. Behavioral findings showed best performance for B faces and, in line with the LVF superiority, better performance for L than R faces. Evoked potentials to B, L, and R faces at 100 to 150-msec poststimulus showed no evidence of asymmetric transfer of information between the hemispheres at early stages of visual processing, suggesting that this factor is not responsible for the LVF superiority. Neural correlates of the LVF superiority, however, were manifested in a shorter latency of the face-specific N170 component to L than R faces and in a larger amplitude to L than R faces at 220—280 and 400—600 msec over both hemispheres. These ERP amplitude differences between L and R faces covaried across subjects with the extent to which the face-specific N170 component was larger over the right than the left hemisphere. We conclude that the two hemispheres exchange information symmetrically at early stages of face processing and together generate a shared facial representation, which is better when facial information is directly presented to the right hemisphere (RH; L faces) than to the left hemisphere (LH; R faces) and best when both hemispheres receive facial information (B faces).

Previous neuropsychological and neuroimaging results have implicated the prefrontal cortex in memory retrieval, although its precise role is unclear. In the present study, we examined patterns of brain electrical activity during retrieval of episodic and semantic memories. In the episodic retrieval task, participants retrieved autobiographical memories in response to event cues. In the semantic retrieval task, participants generated exemplars in response to category cues. Novel sounds presented intermittently during memory retrieval elicited a series of brain potentials including one identifiable as the P3a potential. Based on prior research linking P3a with novelty detection and with the frontal lobes, we predicted that P3a would be reduced to the extent that novelty detection and memory retrieval interfere with each other. Results during episodic and semantic retrieval tasks were compared to results during a task in which subjects attended to the auditory stimuli. P3a amplitudes were reduced during episodic retrieval, particularly at right lateral frontal scalp locations. A similar but less lateralized pattern of frontal P3a reduction was observed during semantic retrieval. These findings support the notion that the right prefrontal cortex is engaged in the service of memory retrieval, particularly for episodic memories.

Many neuropsychological investigations of human memory have focused on the amnesic deficits of alcoholic Korsakoff's syndrome. Structural neuroimaging suggests that the syndrome results from midline diencephalic damage, but functional neuroimaging has the potential to reveal additional neuropathology that may be responsible for cognitive dysfunction. Accordingly, high-resolution positron emission tomography (PET) was used to measure regional cerebral metabolic rates for glucose utilization in five alcoholic Korsakoff patients and nine alcoholic control subjects. Results from a continuous recognition test administered during the radiotracer uptake period indicated that all subjects performed normally with respect to immediate memory, whereas Korsakoff patients demonstrated a marked memory impairment in delayed recognition. PET results from the Korsakoff group showed a widespread decline in glucose metabolism in frontal, parietal, and cingulate regions, suggesting that these functional abnormalities in the cerebral cortex contribute to the memory impairment. Hippocampal glucose metabolism did not differ between the groups. Thus, the evidence did not support the hypothesis that parallel brain dysfunctions are responsible for the similar amnesic symptomatology after hippocampal and diencephalic damage. We hypothesize that the amnesic dysfunction of Korsakoff's syndrome depends on a disruption of thalamocortical interactions that mediate a function critical for normal memory storage.

Event-related brain potentials were recorded from subjects as they attempted to identify words displayed tachistoscopically. Words that had also been presented a few minutes earlier in a different context were identified more often than were words that had not been presented before. This priming effect was observed for words initially seen in an imagery task requiring size estimations as well as for words initially seen in an orthographic task requiring letter counting. Unlike priming, recall and recognition were much better for words repeated from the imagery task than from the orthographic task. Brain potentials elicited during word identification also differed as a function of task. Based on these differences, a potential from 500 to 800 msec was interpreted as an index of recollection processes. Earlier potentials may have indexed processing related to priming. These effects thus provide measures of the hypothetical processes underlying memory performance and demonstrate that recollection and priming are associated with distinct neural events.