Prefrontal cortex and the mediation of proactive interference in working memory

Mediating proactive interference (PI), the deleterious effect of antecedent information on current memory representations, is believed to be a key function of prefrontal cortex (PFC). Item-specific PI results when an invalid probe matches a memorandum from the preceding trial; item-nonspecific PI is produced by the accumulation of no-longer-relevant items from previous trials. We tested the hypothesis that these two types of PI are mediated by common PFC-based processes with an fMRI study of a delayed-recognition task designed to produce both types of PI. Our results indicated that the fMRI correlates of both effects were restricted both to Brodmann’s area 45 in the left hemisphere and to the memory probe epoch of the trial. These results suggest that a unification of the literatures and approaches that have independently studied these phenomena might offer a fruitful new perspective from which to study the relations between working memory, executive control, and the PFC.     

Behavioral and neurophysiological correlates of episodic coding,proactive interference,and list length effects in a running span verbal working memory task

Updating refers to (1) discarding items from, (2) repositioning items in, and (3) adding items to a running working memory span. Our behavioral and fMRI experiments varied three factors: trial length, proactive interference (PI), and group integrity. Group integrity reflected whether the grouping of items at the encoding stage was violated at discarding. Behavioral results were consistent with the idea that updating processes have a relatively short refractory period and may not fatigue, and they revealed that episodic information about group context is encoded automatically in working memory stimulus representations. The fMRI results did not show evidence that updating requirements in a task recruit executive control processes other than those supporting performance on nonupdating trials. They did reveal an item-accumulation effect, in which signal increased monotonically with the number of items presented during the trial, despite the insensitivity of behavioral measures to this factor. Behavioral and fMRI correlates of PI extended previous results and rejected an alternative explanation of PI effects in working memory.     

前摄干扰对工作记忆的作用——大脑如何解决前摄干扰?

前摄干扰(PI)对工作记忆有决定性影响.注意加工理论认为处于工作记忆“注意焦点”的信息不受PI影响,偏向-竞争理论则提出个体对熟悉信息和情景信息代码做出权重评估,解决工作记忆PI效应.研究者从工作记忆编码和提取加工的PI效应、内容相关和情景相关的PI效应、PI消除的脑成像特征等角度进行了验证,多数结果支持偏向-竞争理论.未来研究应使用脑成像技术探讨PI效应与工作记忆子系统、子功能之间的关系等主题.     

Maintenance versus manipulation of information held in working memory: an event-related fMRI study.

One model of the functional organization of lateral prefrontal cortex (PFC) in primates posits that this region is organized in a dorsal/ventral fashion subserving spatial and object working memory, respectively. Alternatively, it has been proposed that a dorsal/ventral subdivision of lateral PFC instead reflects the type of processing performed upon information held in working memory. We tested this hypothesis using an event-related fMRI method that can discriminate among functional changes occurring during temporally separated behavioral subcomponents of a single trial. Subjects performed a delayed-response task with two types of trials in which they were required to: (1) retain a sequence of letters across the delay period (maintenance) or (2) reorder the sequence into alphabetical order across the delay period (manipulation). In each subject, activity during the delay period was found in both dorsolateral and ventrolateral PFC in both types of trials. However, dorsolateral PFC activity was greater in manipulation trials. These findings are consistent with the processing model of the functional organization of working memory in PFC.     

High neuromagnetic activation in the left prefrontal and frontal cortices correlates with better memory performance for abstract words

The ability to store and manipulate online information may be enhanced by an inner speech mechanism that draws upon motor brain regions. Neural correlates of this mechanism were examined using event-related functional magnetic resonance imaging (fMRI). Sixteen participants completed two conditions of a verbal working memory task. In both conditions, participants viewed one or two target letters. In the “storage” condition, these targets were held in mind across a delay. Then a probe letter was presented, and participants indicated by button press whether the probe matched the targets. In the “manipulation” condition, participants identified new targets by thinking two alphabetical letters forward of each original target (e.g., f → h). Participants subsequently indicated whether the probe matched the newly derived targets. Brain activity during the storage and manipulation conditions was examined specifically during the delay phase in order to directly compare manipulation versus storage processes. Activations that were common to both conditions, yet disproportionately greater with manipulation, were observed in the left inferior frontal cortex, premotor cortex, and anterior insula, bilaterally in the parietal lobes and superior cerebellum, and in the right inferior cerebellum. This network shares substrates with overt speech and may represent an inner speech pathway that increases activity with greater working memory demands. Additionally, an inverse correlation was observed between manipulation-related brain activity (on correct trials) and test accuracy in the left premotor cortex, anterior insula, and bilateral superior cerebellum. This inverse relationship may represent intensification of inner speech as one struggles to maintain performance levels.     

Spatial working memory activity of the caudate nucleus is sensitive to frame of reference

We used event-related fMRI to test the hypothesis that the caudate nucleus is preferentially recruited by a spatial working memory task employing egocentrically defined stimuli, which are amenable to transformation into a motor code, as contrasted with allocentrically defined stimuli, which are not. Our results revealed greater delay-epoch activity in egocentric than in allocentric trials in the caudate nucleus and trends in the same direction in the putamen and the lateral premotor cortex (PMC). Response-related activity was greater for egocentric trials in the lateral PMC. We propose that the neostriatum, possibly interacting with the PMC, may contribute to the sensory-motor transformation necessary to establish a prospective motor code (e.g., the representation of a saccade or a grasp). In addition, the PMC may participate in decision-making processes, prompted by the onset of the probe stimulus, that employ this prospective motor information. This model accounts for the empirical evidence that motor distraction disrupts spatial working memory performance.     

Behavioral and neural correlates of memory selection and interference resolution during a digit working memory task

Neuroimaging studies have shown the involvement of prefrontal and posterior parietal cortexes in regulating information processing. We conducted behavioral and fMRI experiments to investigate the relationship between memory selection and proactive interference (PI), using a delayed recognition task with a selection cue presented during the delay indicating which two of the four studied digits were relevant to the present test. PI was indexed by the response time differences between rejecting probes matching and not matching the no longer relevant digits. By varying the delay intervals, we found that the effect of PI did not diminish, even for cases in which the postcue interval was extended to 9 sec, but was stronger when the precue interval was lengthened to 5 sec. By examining the correlation between PI index and neural correlates of memory selection, we found that stronger PI is predicted by lower selection-related activity in the left inferior parietal lobe, the precuneus, and the dorsal middle frontal gyrus. Our results suggest that activity in the prefrontal-parietal network may contribute to one’s ability to focus on the task-relevant information and may proactively reduce PI in working memory.     

The effect of memory load on cortical activity in the spatial working memory circuit

Accumulating evidence from electrophysiology and neuroimaging studies suggests that spatial working memory is subserved by a network of frontal and parietal regions. In the present study, we parametrically varied the memory set size (one to four spatial locations) of a delayed-response task and applied time-resolved fMRI to study the influence of memory load upon the spatial working memory circuit. Our behavioral results showed that performance deteriorates (lower accuracy and longer reaction time) as memory load increases. Memory load influenced cortical activity during the cue, delay, and response phases of the delayed-response task. Although delay-related activity in many regions increased with increasing memory load, it also was significantly reduced in the middle frontal gyrus and frontal eye fields and leveled off in the parietal areas when memory load increased further. Delayrelated activity in the left posterior parietal cortex was also lower during the error trials, in comparison with the correct trials. Our findings indicate that the delay period activity in the spatial working memory circuit is load sensitive and that the attenuation of this signal is the neural manifestation of performance limitation in the face of excessive memory load.     

Contrasting single and multi-component working-memory systems in dual tasking

Working memory can be a major source of interference in dual tasking. However, there is no consensus on whether this interference is the result of a single working memory bottleneck, or of interactions between different working memory components that together form a complete working-memory system. We report a behavioral and an fMRI dataset in which working memory requirements are manipulated during multitasking. We show that a computational cognitive model that assumes a distributed version of working memory accounts for both behavioral and neuroimaging data better than a model that takes a more centralized approach. The model’s working memory consists of an attentional focus, declarative memory, and a subvocalized rehearsal mechanism. Thus, the data and model favor an account where working memory interference in dual tasking is the result of interactions between different resources that together form a working-memory system.     

Individual capacity differences predict working memory performance and prefrontal activity following dopamine receptor stimulation

Dopamine receptors are abundant in the prefrontal cortex (PFC), a critical region involved in working memory. This pharmacological fMRI study tested the relationships between dopamine, PFC function, and individual differences in working memory capacity. Subjects performed a verbal delayed-recognition task after taking either the dopamine receptor agonist bromocriptine or a placebo. Behavioral effects of bromocriptine treatment depended on subjects’ working memory spans, with the greatest behavioral benefit for lower span subjects. After bromocriptine, PFC activity was positively correlated with a measure of cognitive efficiency (RT slope) during the probe period of the task. Less efficient subjects with slower memory retrieval rates had greater PFC activity, whereas more efficient subjects had less activity. After placebo, these measures were uncorrelated. These results support the role of dopamine in verbal working memory and suggest that dopamine may modulate the efficiency of retrieval of items from the contents of working memory. Individual differences in PFC dopamine receptor concentration may thus underlie the behavioral effects of dopamine stimulation on working memory function.     

Effects of chewing on cognitive processing speed

In recent years, chewing has been discussed as producing effects of maintaining and sustaining cognitive performance. We have reported that chewing may improve or recover the process of working memory; however, the mechanisms underlying these phenomena are still to be elucidated. We investigated the effect of chewing on aspects of attention and cognitive processing speed, testing the hypothesis that this effect induces higher cognitive performance. Seventeen healthy adults (20–34 years old) were studied during attention task with blood oxygenation level-dependent functional (fMRI) at 3.0 T MRI. The attentional network test (ANT) within a single task fMRI containing two cue conditions (no cue and center cue) and two target conditions (congruent and incongruent) was conducted to examine the efficiency of alerting and executive control. Participants were instructed to press a button with the right or left thumb according to the direction of a centrally presented arrow. Each participant underwent two back-to-back ANT sessions with or without chewing gum, odorless and tasteless to remove any effect other than chewing. Behavioral results showed that mean reaction time was significantly decreased during chewing condition, regardless of speed-accuracy trade-off, although there were no significant changes in behavioral effects (both alerting and conflict effects). On the other hand, fMRI analysis revealed higher activations in the anterior cingulate cortex and left frontal gyrus for the executive network and motor-related regions for both attentional networks during chewing condition. These results suggested that chewing induced an increase in the arousal level and alertness in addition to an effect on motor control and, as a consequence, these effects could lead to improvements in cognitive performance.     

Motor affordance and its role for visual working memory: evidence from fMRI studies

We examined the role of motor affordances of objects for working memory retention processes. Three experiments are reported in which participants passively viewed pictures of real world objects or had to retain the objects in working memory for a comparison with an S2 stimulus. Brain activation was recorded by means of functional magnetic resonance imaging (fMRI). Retaining information about objects for which hand actions could easily be retrieved (manipulable objects) in working memory activated the hand region of the ventral premotor cortex (PMC) contralateral to the dominant hand. Conversely, nonmanipulable objects activated the left inferior frontal gyrus. This suggests that working memory for objects with motor affordance is based on motor programs associated with their use. An additional study revealed that motor program activation can be modulated by task demands: Holding manipulable objects in working memory for an upcoming motor comparison task was associated with left ventral PMC activation. However, retaining the same objects for a subsequent size comparison task led to activation in posterior brain regions. This suggests that the activation of hand motor programs are under top down control. By this they can flexibly be adapted to various task demands. It is argued that hand motor programs may serve a similar working memory function as speech motor programs for verbalizable working memory contents, and that the premotor system mediates the temporal integration of motor representations with other task-relevant representations in support of goal oriented behavior.     

Trial distinctiveness in visuospatial working memory: effects on individual differences

ABSTRACT

Previous research has shown that proactive interference (PI) negatively affects performance on working memory (WM) tasks, particularly when to-be-remembered items are similar across trials, and that individuals with low WM spans are more susceptible to PI than those with high spans. The two experiments of the present study further explored individual differences in susceptibility to PI in the visuospatial domain. The similarity of the to-be-remembered locations was manipulated across trials by varying whether locations were presented in different colours and/or accompanied by unique sounds. It was hypothesised that these manipulations would reduce PI, improving participants’ memory. In addition, because of their increased susceptibility to PI, it also was hypothesised that low spans would benefit more than high spans from increases in distinctiveness. The results of both experiments were consistent with these hypotheses, suggesting that even low-span individuals can overcome the effects of PI, given the right task support.     

The role of prefrontal cortex in resolving distractor interference

We investigate the hypothesis that those subregions of the prefrontal cortex (PFC) found to support proactive interference resolution may also support delay-spanning distractor interference resolution. Ten subjects performed delayed-recognition tasks requiring working memory for faces or shoes during functional MRI scanning. During the 15-sec delay interval, task-irrelevant distractors were presented. These distractors were either all faces or all shoes and were thus either congruent or incongruent with the domain of items in the working memory task. Delayed-recognition performance was slower and less accurate during congruent than during incongruent trials. Our fMRI analyses revealed significant delay interval activity for face and shoe working memory tasks within both dorsal and ventral PFC. However, only ventral PFC activity was modulated by distractor category, with greater activity for congruent than for incongruent trials. Importantly, this congruency effect was only present for correct trials. In addition to PFC, activity within the fusiform face area was investigated. During face distraction, activity was greater for face relative to shoe working memory. As in ventrolateral PFC, this congruency effect was only present for correct trials. These results suggest that the ventrolateral PFC and fusiform face area may work together to support delay-spanning interference resolution.     

Effects of perceptual quality and visual field of probe stimulus presentation on memory search for letters

Observers indicated whether a single probe letter presented to the left visual field/right hemisphere (LVF-RH) or to the right visual field/left hemisphere (RVF-LH) was contained in a memory set of 2, 3, 4, or 5 letters. For positive trials, the increase in reaction time caused by perceptually degrading the probe letter became progressively larger as memory set size became larger when the probe was presented to the LVF-RH but not when the probe was presented to the RVF-LH. These results were obtained regardless of whether the case of the probe letter varied randomly (Experiment 1) or only capital letters were used (Experiment 2). The results on LVF-RH trials suggest a relatively visuospatial memory comparison process, whereas the results on RVF-LH trials suggest a more abstract memory comparison process. In addition to these effects, the intercept of the memory set size function was lower on LVF-RH trials than on RVF-LH trials when the probe letter was perceptually degraded, consistent with the hypothesis that the right hemisphere is more efficient than the left at early visuospatial processes. Perhaps it is this efficiency at early visuospatial processes that produces the bias toward visuospatial memory comparison on LVF-RH trials.     

Attention on our mind: The role of spatial attention in visual working memory

The current study shows that spatial visual attention is used to retrieve information from visual working memory. Participants had to keep four colored circles in visual working memory. While keeping this information in memory we asked whether one of the colors was present in the array. While retrieving this information, on some trials a probe dot was presented. When this probe dot was presented at the location of the color that had to be retrieved, participants responded faster than when it was presented at another location. Our findings further elaborate the role of visual attention in working memory: not only is attention the mechanism by which information is stored into working memory, it is also the mechanism by which information is retrieved from visual working memory.     

Aging effects on memory encoding in the frontal lobes

Functional magnetic resonance imaging (fMRI) was used to compare frontal-lobe activation in younger and older adults during encoding of words into memory. Participants made semantic or nonsemantic judgments about words. Younger adults exhibited greater activation for semantic relative to nonsemantic judgments in several regions, with the largest activation in the left inferior frontal gyrus. Older adults exhibited greater activation for semantic judgments in the same regions. but the extent of activation was reduced in left prefrontal regions. In older adults, there was a significant association between behavioral tests of declarative and working memory and extent of frontal activation. These results suggest that age-associated decreases in memory ability may be due to decreased frontal-lobe contributions to the initial encoding of experience.     

Proactive interference and item similarity in working memory

Proactive interference (PI) may influence the predictive utility of working memory span tasks. Participants in one experiment (N=70) completed Ravens Advanced Progressive Matrices (RAPM) and multiple versions of operation span and probed recall, modified for the type of memoranda (digits or words). Changing memoranda within- or across-trials released PI, but not doing so permitted PI buildup. Scores from PI-build trials, but not PI-release trials, correlated with RAPM and accounted for as much variance in RAPM as unmodified tasks. These results are consistent with controlled attention and inhibition accounts of working memory, and they elucidate a fundamental component of working memory span tasks.     

Individual differences in proactive interference in verbal and visuospatial working memory

Proactive interference (PI) has been shown to affect working memory (WM) span as well as the predictive utility of WM span measures. However, most of the research on PI has been conducted using verbal memory items, and much less is known about the role of PI in the visuospatial domain. In order to further explore this issue, the present study used a within-subjects manipulation of PI that alternated clusters of trials with verbal and visuospatial to-be-remembered items. Although PI was shown to build and release across trials similarly in the two domains, important differences also were observed. The ability of verbal WM to predict performance on a measure of fluid intelligence was significantly affected by the amount of PI present, consistent with past research, but this proved not to be the case for visuospatial WM. Further, individuals’ susceptibility to PI in one domain was relatively independent of their susceptibility in the other domain, suggesting that, contrary to some theories of executive function, individual differences in PI susceptibility may not be domain-general.     

Are nonlinguistic functions in "Broca's area" prerequisites for language acquisition? FMRI findings from an ontogenetic viewpoint

There is incomplete consensus on the anatomical demarcation of Broca's area in the left inferior frontal gyrus and its functional characterization remains a matter of debate. Exclusive syntactic specialization has been proposed, but is overall inconsistent with the neuroimaging literature. We examined three functional MRI (fMRI) datasets on lexicosemantic decision, tone discrimination, and visuomotor coordination for potential overlap of activation. A single site of convergent activation across all three paradigms was found in the left inferior frontal lobe (area 44/45). This result is discussed in the context of animal and human studies showing inferior frontal participation in visuomotor and audiomotor functions as well as working memory. We propose that Broca's area involvement in lexical semantics and syntax emerges from these nonlinguistic functions, which are prerequisites for language acquisition.