Molecular activity underlying working memory

The prefrontal cortex is necessary for directing thought and planning action. Working memory, the active, transient maintenance of information in mind for subsequent monitoring and manipulation, lies at the core of many simple, as well as high-level, cognitive functions. Working memory has been shown to be compromised in a number of neurological and psychiatric conditions and may contribute to the behavioral and cognitive deficits associated with these disorders. It has been theorized that working memory depends upon reverberating circuits within the prefrontal cortex and other cortical areas. However, recent work indicates that intracellular signals and protein dephosphorylation are critical for working memory. The present article will review recent research into the involvement of the modulatory neurotransmitters and their receptors in working memory. The intracellular signaling pathways activated by these receptors and evidence that indicates a role for G(q)-initiated PI-PLC and calcium-dependent protein phosphatase calcineurin activity in working memory will be discussed. Additionally, the negative influence of calcium- and cAMP-dependent protein kinase (i.e., calcium/calmodulin-dependent protein kinase II (CaMKII), calcium/diacylglycerol-activated protein kinase C (PKC), and cAMP-dependent protein kinase A (PKA)) activities on working memory will be reviewed. The implications of these experimental findings on the observed inverted-U relationship between D(1) receptor stimulation and working memory, as well as age-associated working memory dysfunction, will be presented. Finally, we will discuss considerations for the development of clinical treatments for working memory disorders.     

Recognition and position information in working memory for visual textures

In three experiments, we examined connections between item-recognition memory and memory for item-position information. With sequences of compound gratings as study and probe items, subjects made either item-position judgments (Experiments 1 and 2), by identifying the serial position of the study item that matched the probe, or recognition judgments (Experiment 3), by judging whether the probe had or had not been presented in the study series. Integrating a summed-similarity account of recognition into a signal detection framework shows that the variance of summed similarities on lure trials (probe not present in the study series) exceeds the variance on target trials (probe present in the study series). This prediction is borne out by the empirical zROC functions, all of which had slopes that were greater than 1. Additionally, about 25% of correct recognitions were accompanied by incorrect item position identification. Misidentifications of item position arose from two sources--structural similarity and positional similarity-which combined in an approximately additive fashion.     

Visual working memory for global,object, and part-based information

We investigated visual working memory for novel objects and parts of novel objects. After a delay period, participants showed strikingly more accurate performance recognizing a single whole object than the parts of that object. This bias to remember whole objects, rather than parts, persisted even when the division between parts was clearly defined and the parts were disconnected from each other so that, in order to remember the single whole object, the participants needed to mentally combine the parts. In addition, the bias was confirmed when the parts were divided by color. These experiments indicated that holistic perceptual-grouping biases are automatically used to organize storage in visual working memory. In addition, our results suggested that the bias was impervious to top-down consciously directed control, because when task demands were manipulated through instruction and catch trials, the participants still recognized whole objects more quickly and more accurately than their parts. This bias persisted even when the whole objects were novel and the parts were familiar. We propose that visual working memory representations depend primarily on the global configural properties of whole objects, rather than part-based representations, even when the parts themselves can be clearly perceived as individual objects. This global configural bias beneficially reduces memory load on a capacity-limited system operating in a complex visual environment, because fewer distinct items must be remembered.     

Guidance of attention by information held in working memory

Information held in working memory (WM) can guide attention during visual search. The authors of recent studies have interpreted the effect of holding verbal labels in WM as guidance of visual attention by semantic information. In a series of experiments, we tested how attention is influenced by visual features versus category-level information about complex objects held in WM. Participants either memorized an object’s image or its category. While holding this information in memory, they searched for a target in a four-object search display. On exact-match trials, the memorized item reappeared as a distractor in the search display. On category-match trials, another exemplar of the memorized item appeared as a distractor. On neutral trials, none of the distractors were related to the memorized object. We found attentional guidance in visual search on both exact-match and category-match trials in Experiment 1, in which the exemplars were visually similar. When we controlled for visual similarity among the exemplars by using four possible exemplars (Exp. 2) or by using two exemplars rated as being visually dissimilar (Exp. 3), we found attentional guidance only on exact-match trials when participants memorized the object’s image. The same pattern of results held when the target was invariant (Exps. 2–3) and when the target was defined semantically and varied in visual features (Exp. 4). The findings of these experiments suggest that attentional guidance by WM requires active visual information.     

Updating information in verbal working memory and executive functioning

50 older adults (M age=77.9 yr., SD=7.3; 35 women and 15 men) were tested using the updating working-memory task. They were also given the neuropsychological Wisconsin Card Sorting Test, assumed to evaluate executive functioning and the frontal cortex. A factor analysis with age, education, and verbal ability partialled out was computed on the updating task outcomes and resulted in a two-factor solution, indicating that this task requires two independent processes, interpreted as reflecting a storage component and an updating component. Partial correlations with age, education, and verbal ability partialled out indicated that Wisconsin Card Sorting Test measures were significantly associated with the factor supposed to reflect the updating process. Such results appeared consistent with the model of working memory with a central executive system involved in the updating process and related to the executive-frontal functioning, and a phonological loop system involved in the storage of verbal information and not linked to executive-frontal functions.     

Sensitive maintenance: a cognitive process underlying individual differences in memory for threatening information

Dispositional styles of coping with threat influence memory for threatening information. In particular, sensitizers excel over repressors in their memory for threatening information after long retention intervals, but not after short ones. We therefore suggested that sensitizers, but not repressors, employ active maintenance processes during the retention interval to selectively retain threatening material. Sensitive maintenance was studied in 2 experiments in which participants were briefly exposed to threatening and nonthreatening pictures (Experiment 1, N = 128) or words (Experiment 2, N = 145). Following, we administered unannounced recognition tests before and after an intervening task that generated either high or low cognitive load, assuming that high cognitive load would impede sensitizers' memory maintenance of threatening material. Supporting our hypotheses, the same pattern of results was obtained in both experiments: Under low cognitive load, sensitizers forgot less threat material than repressors did; no such differences were observed under high cognitive load.     

Consolidation of statistical information of multiple objects in working memory

The present study investigated working memory consolidation in focused and distributed attention tasks by examining the time course of the consolidation process (Experiment 1) and its dependence on capacity-limited central resources (Experiment 2) in both tasks. In a match-to-sample design using masks at various intervals to vary consolidation rates, the participants performed either an identification task (focused attention) or a mean estimation task (distributed attention) with (Experiment 1) or without (Experiment 2) prior knowledge of what task they were to perform. We found that consolidation in the distributed attention task was more efficient and was about twice as fast as in the focused attention task. In addition, both tasks suffered interference when they had to be performed together, indicating that both types of attention rely on a common set of control processes. These findings can be attributed to differences in the resolution of object representations and in the scope of attention associated with focused and distributed attention.     

The neural correlates of updating information in verbal working memory

The aim of the present study was to re-examine cerebral areas subserving the updating function of the central executive with a running span task requiring subjects to watch strings of consonants of unknown length and then to recall serially a specific number of recent items. In order to dissociate more precisely the updating process from the storage function, a four-item instead of a six-item memory load was used, contrary to our previous study (Salmon et al., 1996). In addition, a serial recall procedure was preferred to a recognition procedure in order to suppress the use of visuospatial strategies. The most significant increase of rCBF occurred in the left frontopolar cortex (Brodmann's area 10), spreading to the left middle frontal (Brodmann's area 46). Results suggest that frontopolar activation underlies an updating process in working memory.     

Misleading postevent information and working memory capacity: an individual differences approach to eyewitness memory

The present study examines how individual differences in working memory capacity relate to the effect of misleading postevent information on memory for the original event. Participants were shown a film of a crime event and were then asked to unscramble a narrative that included misinformation regarding some of the film's details. Additionally, the working memory capacity of the participants was measured using the operation‐word span task. Finally, in a free recall test, participants recalled fewer correct details in the misinformation condition compared to the control condition. This effect was negatively correlated with working memory capacity. Copyright © 2002 John Wiley & Sons, Ltd.     

Visual working memory for observed actions

Human society depends on the ability to remember the actions of other individuals, which is information that must be stored in a temporary buffer to guide behavior after actions have been observed. To date, however, the storage capacity, contents, and architecture of working memory for observed actions are unknown. In this article, the author shows that it is possible to retain information about only 2-3 actions in visual working memory at once. However, it is also possible to retain 9 properties distributed across 3 actions almost as well as 3 properties distributed across 3 actions, showing that working memory stores integrated action representations rather than individual properties. Finally, the author shows that working memory for observed actions is independent from working memory for object and spatial information. These results provide evidence for a previously undocumented system in working memory for storing information about actions. Further, this system operates by the same storage principles as visual working memory for object information. Thus, working memory consists of a series of distinct yet computationally similar mechanisms for retaining different types of visual information.     

Distraction biases working memory for faces

Psychonomic Bulletin & Review - Working memory persists in the face of distraction, yet not without consequence. Previous research has shown that memory for low-level visual features is...     

Implicit working memory

Working Memory (WM) plays a crucial role in many high-level cognitive processes (e.g., reasoning, decision making, goal pursuit and cognitive control). The prevalent view holds that active components of WM are predominantly intentional and conscious. This conception is oftentimes expressed explicitly, but it is best reflected in the nature of major WM tasks: All of them are blatantly explicit. We developed two new WM paradigms that allow for an examination of the role of conscious awareness in WM. Results from five studies show that WM can operate unintentionally and outside of conscious awareness, thus suggesting that the current view should be expanded to include implicit WM.     

Dissociation of the neural systems for working memory maintenance of verbal and nonspatial visual information

Working memory for names and faces was investigated to ascertain whether verbal and nonspatial visual information is maintained in working memory by separate neural systems. The subjects performed a delayed match-to-sample task for famous or unfamous faces and names and a sensorimotor control task. Several occipital, temporal, parietal, and prefrontal areas were activated during all memory delays, in comparison with the control delays. Greater delay activity for unfamous faces than for names was obtained in the right fusiform gyrus, right inferior frontal gyrus (IFG), right IFG/ precentral gyrus, and right medial superior frontal gyrus, whereas greater delay activity for unfamous names than for faces was observed in the precuneus, left insula/postcentral gyrus, and left IFG/ precentral gyrus. There was no significant difference in the prefrontal activity in the comparison between famous faces and names. Greater delay activity for famous names than for faces was obtained in visual association and parietal areas. The results indicate that there is a functional dissociation based on information type within the neural system that is responsible for working memory maintenance of verbal and nonspatial visual information.     

Access to information in working memory: exploring the focus of attention

Participants memorized briefly presented sets of digits, a subset of which had to be accessed as input for arithmetic tasks (the active set), whereas another subset had to be remembered independently of the concurrent task (the passive set). Latencies for arithmetic operations were a function of the setsize of active but not passive sets. Object-switch costs were observed when successive operations were applied to different digits within an active set. Participants took 2 s to encode a passive set so that it did not affect processing latencies (Experiment 2). The results support a model distinguishing 3 states of representations in working memory: the activated part of long-term memory, a capacity limited region of direct access, and a focus of attention.     

Processing of temporal duration information in working memory after frontodorsal tumour excisions

This study aimed to test the hypothesis that impairments of temporal duration processing after frontal lobe lesions reflect deficits in executive monitoring functions rather than a domain-specific deficit in the maintenance of duration information in working memory. Patients with frontodorsal lesions, clinical controls with post-central lesions, and healthy controls performed recognition and classification tasks, which should allow for testing maintenance and monitoring functions, respectively. Results showed mild non-selective impairments of the frontal patients on both temporal and spatial recognition tasks, but a marked selective degradation on temporal classification while performance on spatial classification was unimpaired. This suggests that maintenance of duration information in working memory after frontal lesions is basically preserved but that, depending on executive task characteristics, there is a specific deficit in the strategic organization of this type of information.     

Effect of normal aging on the manipulation of information in working memory

The goal of this study is to examine the central executive of working memory in normal aging, specifically focusing on its capacities to manipulate or modify the format of the to-be-recalled material. The central executive was measured with the alphabetical span procedure, during which subjects were asked to recall a random series of words in their alphabetical order. The storage demand was equalized across subjects by adjusting the list lengths according to individual span. Experiments 1, 2, and 3 showed that elderly subjects were not impaired in manipulating information, relative to young controls, even when the difficulty of the task was increased. In Experiment 4, validity was tested by asking young subjects to perform the task under the conditions of full or divided attention. Alphabetical recall was more impaired than direct recall during the divided attention condition, which suggests a larger involvement of the central executive component in the former. These results are discussed in relation to the hypothesis of a central executive impairment being associated with normal aging.     

Distinguishing short-term memory from working memory

The aim of the present research was to determine whether short-term memory and working memory could be distinguished. In two studies, 7- to 13-year-olds (N = 155, N = 132) were administered tasks thought to assess short-term memory as well as tasks thought to assess working memory. Both exploratory and confirmatory factor analyses distinguished short-term memory tasks from working memory tasks. In addition, performance on working memory tasks was related to word decoding skill but performance on short-term memory tasks was not. Finally, performance on both short-term memory and working memory tasks were associated with age-related increases in processing speed. Results are discussed in relation to models of short-term and working memory.     

Mechanisms underlying the slant aftereffect

Transfer of the median plane slant aftereffect was assessed across changes in the type of depth information for the slant of the display. In addition, the effectiveness of monocularpictorial and binocular information in inducing the aftereffect was measured. Binocular information produced a larger aftereffect than did monocular-pictorial information, and adaptation created with one type of depth information induced an aftereffect assessed with presentation of the other type of depth information. The results suggest that the slant aftereffect is not entirely specific to type of depth information presented. The induction of the aftereffect involves a process more general than the sensory mechanisms responsible for adaptation to twodimensional tilt or adaptation to a texture gradient.     

Preserved learning of novel information in amnesia: Evidence for multiple memory systems

Four of five patients with marked global amnesia, and others with new learning impairments, showed normal processing facilitation for novel stimuli (nonwords) and/or for familiar stimuli (words) on a word/nonword (lexical) decision task. The data are interpreted as a reflection of the learning capabilities of in-line neural processing stages with multiple, distinct, informational codes. These in-line learning processes are separate from the recognition/recall memory impaired by amygdalohippocampal/dosomedial thalamic damage, but probably supplement such memory in some tasks in normal individuals. Preserved learning of novel information seems incompatible with explanations of spared learning in amnesia that are based on the episodic/semantic or memory/habit distinctions, but is consistent with the procedural/declarative hypothesis.     

Precision of working memory for speech sounds

Memory for speech sounds is a key component of models of verbal working memory (WM). But how good is verbal WM? Most investigations assess this using binary report measures to derive a fixed number of items that can be stored. However, recent findings in visual WM have challenged such “quantized” views by employing measures of recall precision with an analogue response scale. WM for speech sounds might rely on both continuous and categorical storage mechanisms. Using a novel speech matching paradigm, we measured WM recall precision for phonemes. Vowel qualities were sampled from a formant space continuum. A probe vowel had to be adjusted to match the vowel quality of a target on a continuous, analogue response scale. Crucially, this provided an index of the variability of a memory representation around its true value and thus allowed us to estimate how memories were distorted from the original sounds. Memory load affected the quality of speech sound recall in two ways. First, there was a gradual decline in recall precision with increasing number of items, consistent with the view that WM representations of speech sounds become noisier with an increase in the number of items held in memory, just as for vision. Based on multidimensional scaling (MDS), the level of noise appeared to be reflected in distortions of the formant space. Second, as memory load increased, there was evidence of greater clustering of participants' responses around particular vowels. A mixture model captured both continuous and categorical responses, demonstrating a shift from continuous to categorical memory with increasing WM load. This suggests that direct acoustic storage can be used for single items, but when more items must be stored, categorical representations must be used.