Episodic memory formation in unrestricted viewing.

The brain systems of episodic memory and oculomotor control are tightly linked, suggesting a crucial role of eye movements in memory. But little is known about the neural mechanisms of memory formation across eye movements in unrestricted viewing behavior. Here, we leverage simultaneous eye tracking and EEG recording to examine episodic memory formation in free viewing. Participants memorized multi-element events while their EEG and eye movements were concurrently recorded. Each event comprised elements from three categories (face, object, place), with two exemplars from each category, in different locations on the screen. A subsequent associative memory test assessed participants' memory for the between-category associations that specified each event. We used a deconvolution approach to overcome the problem of overlapping EEG responses to sequential saccades in free viewing. Brain activity was time-locked to the fixation onsets, and we examined EEG power in the theta and alpha frequency bands, the putative oscillatory correlates of episodic encoding mechanisms. Three modulations of fixation-related EEG predicted high subsequent memory performance: (1) theta increase at fixations after between-category gaze transitions, (2) theta and alpha increase at fixations after within-element gaze transitions, (3) alpha decrease at fixations after between-exemplar gaze transitions. Thus, event encoding with unrestricted viewing behavior was characterized by three neural mechanisms, manifested in fixation-locked theta and alpha EEG activity that rapidly turned on and off during the unfolding eye movement sequences. These three distinct neural mechanisms may be the essential building blocks that subserve the buildup of coherent episodic memories during unrestricted viewing behavior.

Encoding contexts are incidentally reinstated during competitive retrieval and track the temporal dynamics of memory interference.

The ability to remember an episode from our past is often hindered by competition from similar events. For example, if we want to remember the article a colleague recommended during the last lab meeting, we may need to resolve interference from other article recommendations from the same colleague. This study investigates if the contextual features specifying the encoding episodes are incidentally reinstated during competitive memory retrieval. Competition between memories was created through the AB/AC interference paradigm. Individual word-pairs were presented embedded in a slowly drifting real-word-like context. Multivariate pattern analysis (MVPA) of high temporal-resolution electroencephalographic (EEG) data was used to investigate context reactivation during memory retrieval. Behaviorally, we observed proactive (but not retroactive) interference; that is, performance for AC competitive retrieval was worse compared with a control DE noncompetitive retrieval, whereas AB retrieval did not suffer from competition. Neurally, proactive interference was accompanied by an early reinstatement of the competitor context and interference resolution was associated with the ensuing reinstatement of the target context. Together, these findings provide novel evidence showing that the encoding contexts of competing discrete events are incidentally reinstated during competitive retrieval and that such reinstatement tracks retrieval competition and subsequent interference resolution.

Strategic retrieval prevents memory interference: The temporal dynamics of retrieval orientation.

Resolving interference between overlapping memories is crucial to remember the past. This study tests the novel prediction that orienting search focus benefits goal-relevant retrieval by reducing competition from unwanted memories. In a modified retrieval-practice paradigm, participants encoded word-pairs in one of two encoding tasks. Critically, to evaluate whether this retrieval orientation (RO) reduces memory interference, target and competitor memories were always related to different encoding tasks. At retrieval, instructions were provided for half of the blocks with the intention to bias remembering towards items encoded with one of the ROs. Behavioural data show that adopting an RO improved target accessibility, strengthened the testing effect, and reduced retrieval-induced forgetting (RIF) of competitors. Specifically, RIF - typically attributed to inhibitory control of memory interference - was prominent when no retrieval orientation (NRO) instruction was provided. Furthermore, a neural correlate of RO was calculated by training a linear discriminant analysis (LDA) to discriminate the electroencephalographic (EEG) spatial brain patterns correspondent to the two ROs over the time course of selective retrieval. RO was characterised by increases in the theta and decreases in the beta frequency band, evident both before and after category-cue onset. While the pre-cue RO reinstatement effect predicted both immediate retrieval-practice success and later target accessibility, the post-cue effect predicted disengagement of inhibitory control, such that participants showing a stronger RO reinstatement effect showed lower levels of RIF. These data suggest that strategically orienting search focus during retrieval both increases target memory accessibility and reduces memory interference, which consequently protects related memories from inhibition and later forgetting. Furthermore, they also highlight the roles of theta and beta oscillations in establishing and maintaining a task-relevant bias towards target memory representations during competitive memory retrieval.

Temporal Dynamics of Memory-guided Cognitive Control and Generalization of Control via Overlapping Associative Memories.

Goal-directed behavior can benefit from proactive adjustments of cognitive control that occur in anticipation of forthcoming cognitive control demands (CCD). Predictions of forthcoming CCD are thought to depend on learning and memory in two ways: First, through direct experience, associative encoding may link previously experienced CCD to its triggering item, such that subsequent encounters with the item serve to cue retrieval of (i.e., predict) the associated CCD. Second, in the absence of direct experience, pattern completion and mnemonic integration mechanisms may allow CCD to be generalized from its associated item to other items related in memory. While extant behavioral evidence documents both types of CCD prediction, the neurocognitive mechanisms giving rise to these predictions remain largely unexplored. Here, we tested two hypotheses: (1) memory-guided predictions about CCD precede control adjustments due to the actual CCD required; and (2) generalization of CCD can be accomplished through integration mechanisms that link partially overlapping CCD-item and item-item associations in memory. Supporting these hypotheses, the temporal dynamics of theta and alpha power in human electroencephalography data (n = 43, 26 females) revealed that an associative CCD effect emerges earlier than interaction effects involving actual CCD. Furthermore, generalization of CCD from one item (X) to another item (Y) was predicted by a decrease in alpha power following the presentation of the X-Y pair. These findings advance understanding of the mechanisms underlying memory-guided adjustments of cognitive control.SIGNIFICANCE STATEMENT Cognitive control adaptively regulates information processing to align with task goals. Experience-based expectations enable adjustments of control, leading to improved performance when expectations match the actual control demand required. Using EEG, we demonstrate that memory for past cognitive control demand proactively guides the allocation of cognitive control, preceding adjustments of control triggered by the demands of the present environment. Furthermore, we demonstrate that learned cognitive control demands can be generalized through mnemonic integration processes, enabling the spread of expectations about cognitive control demands to items associated in memory. We reveal that this generalization is linked to decreased alpha oscillation in medial frontal channels. Collectively, these findings provide new insights into how memory-control interactions facilitate goal-directed behavior.

Neural Pattern Classification Tracks Transfer-Appropriate Processing in Episodic Memory.

The transfer-appropriate processing (TAP) account holds that episodic memory depends on the overlap between encoding and retrieval processing. In the current study, we employed multivariate pattern analysis (MVPA) of electroencephalography to examine the relevance of spontaneously engaged visual processing during encoding for later retrieval. Human participants encoded word-picture associations, where the picture could be a famous face, a landmark, or an object. At test, we manipulated the retrieval demands by asking participants to retrieve either visual or verbal information about the pictures. MVPA revealed classification between picture categories during early perceptual stages of encoding (∼170 ms). Importantly, these visual category-specific neural patterns were predictive of later episodic remembering, but the direction of the relationship was contingent on the particular retrieval demand of the memory task: a benefit for the visual and a cost for the verbal. A reinstatement of the category-specific neural patterns established during encoding was observed during retrieval, and again the relationship with behavior varied with retrieval demands. Reactivation of visual representations during retrieval was associated with better memory in the visual task, but with lower performance in the verbal task. Our findings support and extend the TAP account by demonstrating that processing of particular aspects during memory formation can also have detrimental effects on later episodic remembering when other aspects of the event are called-for and shed new light on encoding and retrieval interactions in episodic memory.

Implicit sequence learning is preserved in dyslexic children.

This study investigates the implicit sequence learning abilities of dyslexic children using an artificial grammar learning task with an extended exposure period. Twenty children with developmental dyslexia participated in the study and were matched with two control groups-one matched for age and other for reading skills. During 3 days, all participants performed an acquisition task, where they were exposed to colored geometrical forms sequences with an underlying grammatical structure. On the last day, after the acquisition task, participants were tested in a grammaticality classification task. Implicit sequence learning was present in dyslexic children, as well as in both control groups, and no differences between groups were observed. These results suggest that implicit learning deficits per se cannot explain the characteristic reading difficulties of the dyslexics.

Mental reinstatement of encoding context improves episodic remembering.

This study investigates context-dependent memory retrieval. Previous work has shown that physically re-experiencing the encoding context at retrieval improves memory accessibility. The current study examined if mental reconstruction of the original encoding context would yield parallel memory benefits. Participants performed a cued-recall memory task, preceded either by a mental or by a physical context reinstatement task, and we manipulated whether the context reinstated at retrieval overlapped with the context of the target episode. Both behavioral and electrophysiological measures of brain activity showed strong encoding-retrieval (E-R) overlap effects, with facilitated episodic retrieval when the encoding and retrieval contexts overlapped. The electrophysiological E-R overlap effect was more sustained and involved more posterior regions when context was mentally compared with physically reinstated. Additionally, a time-frequency analysis revealed that context reinstatement alone engenders recollection of the target episode. However, while recollection of the target memory is readily prompted by a physical reinstatement, target recollection during mental reinstatement is delayed and depends on the gradual reconstruction of the context. Taken together, our results show facilitated episodic remembering also when mentally reinstating the encoding context; and that such benefits are supported by both shared and partially non-overlapping neural mechanisms when the encoding context is mentally reconstructed as compared with physically presented at the time of retrieval.

Benefits and Costs of Context Reinstatement in Episodic Memory: An ERP Study.

This study investigated context-dependent episodic memory retrieval. An influential idea in the memory literature is that performance benefits when the retrieval context overlaps with the original encoding context. However, such memory facilitation may not be driven by the encoding-retrieval overlap per se but by the presence of diagnostic features in the reinstated context that discriminate the target episode from competing episodes. To test this prediction, the encoding-retrieval overlap and the diagnostic value of the context were manipulated in a novel associative recognition memory task. Participants were asked to memorize word pairs presented together with diagnostic (unique) and nondiagnostic (shared) background scenes. At test, participants recognized the word pairs in the presence and absence of the previously encoded contexts. Behavioral data show facilitated memory performance in the presence of the original context but, importantly, only when the context was diagnostic of the target episode. The electrophysiological data reveal an early anterior ERP encoding-retrieval overlap effect that tracks the cost associated with having nondiagnostic contexts present at retrieval, that is, shared by multiple previous episodes, and a later posterior encoding-retrieval overlap effect that reflects facilitated access to the target episode during retrieval in diagnostic contexts. Taken together, our results underscore the importance of the diagnostic value of the context and suggest that context-dependent episodic memory effects are multiple determined.

Lexical and sublexical orthographic processing: an ERP study with skilled and dyslexic adult readers.

This ERP study investigated the cognitive nature of the P1-N1 components during orthographic processing. We used an implicit reading task with various types of stimuli involving different amounts of sublexical or lexical orthographic processing (words, pseudohomophones, pseudowords, nonwords, and symbols), and tested average and dyslexic readers. An orthographic regularity effect (pseudowords-nonwords contrast) was observed in the average but not in the dyslexic group. This suggests an early sensitivity to the dependencies among letters in word-forms that reflect orthographic structure, while the dyslexic brain apparently fails to be appropriately sensitive to these complex features. Moreover, in the adults the N1-response may already reflect lexical access: (i) the N1 was sensitive to the familiar vs. less familiar orthographic sequence contrast; (ii) and early effects of the phonological form (words-pseudohomophones contrast) were also found. Finally, the later N320 component was attenuated in the dyslexics, suggesting suboptimal processing in later stages of phonological analysis.

Lexical and phonological processes in dyslexic readers: evidence from a visual lexical decision task.

The aim of the present study was to investigate whether reading failure in the context of an orthography of intermediate consistency is linked to inefficient use of the lexical orthographic reading procedure. The performance of typically developing and dyslexic Portuguese-speaking children was examined in a lexical decision task, where the stimulus lexicality, word frequency and length were manipulated. Both lexicality and length effects were larger in the dyslexic group than in controls, although the interaction between group and frequency disappeared when the data were transformed to control for general performance factors. Children with dyslexia were influenced in lexical decision making by the stimulus length of words and pseudowords, whereas age-matched controls were influenced by the length of pseudowords only. These findings suggest that non-impaired readers rely mainly on lexical orthographic information, but children with dyslexia preferentially use the phonological decoding procedure--albeit poorly--most likely because they struggle to process orthographic inputs as a whole such as controls do. Accordingly, dyslexic children showed significantly poorer performance than controls for all types of stimuli, including words that could be considered over-learned, such as high-frequency words. This suggests that their orthographic lexical entries are less established in the orthographic lexicon.

Electrophysiological correlates of impaired reading in dyslexic pre-adolescent children.

In this study, event related potentials (ERPs) were used to investigate the extent to which dyslexics (aged 9-13 years) differ from normally reading controls in early ERPs, which reflect prelexical orthographic processing, and in late ERPs, which reflect implicit phonological processing. The participants performed an implicit reading task, which was manipulated in terms of letter-specific processing, orthographic familiarity, and phonological structure. Comparing consonant- and symbol sequences, the results showed significant differences in the P1 and N1 waveforms in the control but not in the dyslexic group. The reduced P1 and N1 effects in pre-adolescent children with dyslexia suggest a lack of visual specialization for letter-processing. The P1 and N1 components were not sensitive to the familiar vs. less familiar orthographic sequence contrast. The amplitude of the later N320 component was larger for phonologically legal (pseudowords) compared to illegal (consonant sequences) items in both controls and dyslexics. However, the topographic differences showed that the controls were more left-lateralized than the dyslexics. We suggest that the development of the mechanisms that support literacy skills in dyslexics is both delayed and follows a non-normal developmental path. This contributes to the hemispheric differences observed and might reflect a compensatory mechanism in dyslexics.

The interaction between surface color and color knowledge: behavioral and electrophysiological evidence.

In this study, we used event-related potentials (ERPs) to evaluate the contribution of surface color and color knowledge information in object identification. We constructed two color-object verification tasks - a surface and a knowledge verification task - using high color diagnostic objects; both typical and atypical color versions of the same object were presented. Continuous electroencephalogram was recorded from 26 subjects. A cluster randomization procedure was used to explore the differences between typical and atypical color objects in each task. In the color knowledge task, we found two significant clusters that were consistent with the N350 and late positive complex (LPC) effects. Atypical color objects elicited more negative ERPs compared to typical color objects. The color effect found in the N350 time window suggests that surface color is an important cue that facilitates the selection of a stored object representation from long-term memory. Moreover, the observed LPC effect suggests that surface color activates associated semantic knowledge about the object, including color knowledge representations. We did not find any significant differences between typical and atypical color objects in the surface color verification task, which indicates that there is little contribution of color knowledge to resolve the surface color verification. Our main results suggest that surface color is an important visual cue that triggers color knowledge, thereby facilitating object identification.

Object naming in dyslexic children: more than a phonological deficit.

In the present study, the authors investigate how some visual factors related to early stages of visual-object naming modulate naming performance in dyslexia. The performance of dyslexic children was compared with 2 control groups-normal readers matched for age and normal readers matched for reading level-while performing a discrete naming task in which color and dimensionality of the visually presented objects were manipulated. The results showed that 2-dimensional naming performance improved for color representations in control readers but not in dyslexics. In contrast to control readers, dyslexics were also insensitive to the stimulus's dimensionality. These findings are unlikely to be explained by a phonological processing problem related to phonological access or retrieval but suggest that dyslexics have a lower capacity for coding and decoding visual surface features of 2-dimensional representations or problems with the integration of visual information stored in long-term memory.

The role of color information on object recognition: a review and meta-analysis.

In this study, we systematically review the scientific literature on the effect of color on object recognition. Thirty-five independent experiments, comprising 1535 participants, were included in a meta-analysis. We found a moderate effect of color on object recognition (d=0.28). Specific effects of moderator variables were analyzed and we found that color diagnosticity is the factor with the greatest moderator effect on the influence of color in object recognition; studies using color diagnostic objects showed a significant color effect (d=0.43), whereas a marginal color effect was found in studies that used non-color diagnostic objects (d=0.18). The present study did not permit the drawing of specific conclusions about the moderator effect of the object recognition task; while the meta-analytic review showed that color information improves object recognition mainly in studies using naming tasks (d=0.36), the literature review revealed a large body of evidence showing positive effects of color information on object recognition in studies using a large variety of visual recognition tasks. We also found that color is important for the ability to recognize artifacts and natural objects, to recognize objects presented as types (line-drawings) or as tokens (photographs), and to recognize objects that are presented without surface details, such as texture or shadow. Taken together, the results of the meta-analysis strongly support the contention that color plays a role in object recognition. This suggests that the role of color should be taken into account in models of visual object recognition.

The influence of color information on the recognition of color diagnostic and noncolor diagnostic objects.

In the present study, the authors explore in detail the level of visual object recognition at which perceptual color information improves the recognition of color diagnostic and noncolor diagnostic objects. To address this issue, 3 object recognition tasks with different cognitive demands were designed: (a) an object verification task; (b) a category verification task; and (c) a name verification task. The authors found that perceptual color information improved color diagnostic object recognition mainly in tasks for which access to the semantic knowledge about the object was necessary to perform the task; that is, in category and name verification. In contrast, the authors found that perceptual color information facilitates noncolor diagnostic object recognition when access to the object's structural description from long-term memory was necessary--that is, object verification. In summary, the present study shows that the role of perceptual color information in object recognition is dependent on color diagnosticity.

Cortical brain regions associated with color processing: an FMRI study.

To clarify whether the neural pathways concerning color processing are the same for natural objects, for artifacts objects and for non-objects we examined brain responses measured with functional magnetic resonance imaging (FMRI) during a covert naming task including the factors color (color vs. black&white (B&W)) and stimulus type (natural vs. artifacts vs. non-objects). Our results indicate that the superior parietal lobule and precuneus (BA 7) bilaterally, the right hippocampus and the right fusifom gyrus (V4) make part of a network responsible for color processing both for natural objects and artifacts, but not for non-objects. When color objects (both natural and artifacts) were contrasted with color non-objects we observed activations in the right parahippocampal gyrus (BA 35/36), the superior parietal lobule (BA 7) bilaterally, the left inferior middle temporal region (BA 20/21) and the inferior and superior frontal regions (BA 10/11/47). These additional activations suggest that colored objects recruit brain regions that are related to visual semantic information/retrieval and brain regions related to visuo-spatial processing. Overall, the results suggest that color information is an attribute that can improve object recognition (behavioral results) and activate a specific neural network related to visual semantic information that is more extensive than for B&W objects during object recognition.

The influence of surface color information and color knowledge information in object recognition.

In order to clarify whether the influence of color knowledge information in object recognition depends on the presence of the appropriate surface color, we designed a name-object verification task. The relationship between color and shape information provided by the name and by the object photo was manipulated in order to assess color interference independently of shape interference. We tested three different versions for each object: typically colored, black and white, and nontypically colored. The response times on the nonmatching trials were used to measure the interference between the name and the photo. We predicted that the more similar the name and the photo are, the longer it would take to respond. Overall, the color similarity effect disappeared in the black-and-white and nontypical color conditions, suggesting that the influence of color knowledge on object recognition depends on the presence of the appropriate surface color information.

The impact of reading and writing skills on a visuo-motor integration task: a comparison between illiterate and literate subjects.

Previous studies have shown a significant association between reading skills and the performance on visuo-motor tasks. In order to clarify whether reading and writing skills modulate non-linguistic domains, we investigated the performance of two literacy groups on a visuo-motor integration task with non-linguistic stimuli. Twenty-one illiterate participants and twenty matched literate controls were included in the experiment. Subjects were instructed to use the right or the left index finger to point to and touch a randomly presented target on the right or left side of a touch screen. The results showed that the literate subjects were significantly faster in detecting and touching targets on the left compared to the right side of the screen. In contrast, the presentation side did not affect the performance of the illiterate group. These results lend support to the idea that having acquired reading and writing skills, and thus a preferred left-to-right reading direction, influences visual scanning.