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.     

Those cheating rats: male and female rats use odor trails in a water-escape "working memory" task.

Three-month-old Sprague-Dawley rats were trained on a working memory win-stay (spatial delayed matching-to-sample) water-escape task with the escape platform location the same for all subjects on a given trial, a procedure that maximizes the buildup of an odor trail to the escape platform. In subsequent tests during which the location of the escape platform varied randomly between subjects, the rats, especially the females, while continuing to perform above chance level, made increased errors. Varying the platform location between subjects eliminated odor trail as a nonambiguous cue for locating the escape platform. In a second experiment females performed better than males on a reference memory odor trail discrimination task which involved following the path of like-gender "pathmaker" rats to the escape platform. The relatively poor use of odor trails by the males was associated with a high frequency of choosing a preferred choice section or returning to the choice section selected first on the immediately preceding trial (perseveration). Collectively, the two experiments demonstrate that rats can use either working memory or odor trails to locate an escape platform in a water maze, and that they, especially females, will use odor trails in a working memory task if odor trails are available. Clearly, the location of the escape platform should be varied randomly between subjects in tests of working memory.     

The role of response and reward in spatial memory

Two experiments were conducted to determine the role of the response and of reward in spatial working memory. Rats were initially trained on a four-arm maze to run to the end of each arm for a single pellet of food. On subsequent tests, rats were first placed at the end of one, two, or three arms. In Experiment 1, the arms on which the rat was placed (“placed arms”) had food which the rat was allowed to eat, whereas in Experiment 2 these placed arms did not have food. Following the placements the rat was allowed to choose among the four arms; only unplaced arms contained food. Two measures indicated that the response made a slight but reliable contribution to spatial memory. (a) When a rewarded arm was still available, choice accuracy after placements was less than choice accuracy on tests in which no placements had occurred; this difference diminished over test days. (b) When all four arms had been chosen once, the rats were more likely to go back to a placed arm rather than an unplaced arm. No influence of the presence or absence of food on the placed arms was found. These data demonstrate that the response of running down an arm, but not the reward outcome at the end, had a small influence on the memorability of a visit. Overall, above chance performance in the spatial working memory task was maintained without either running to the arm or obtaining food on it.     

Effects of verbal and nonverbal interference on spatial and object visual working memory

We tested the hypothesis that a verbal coding mechanism is necessarily engaged by object, but not spatial, visual working memory tasks. We employed a dual-task procedure that paired n-back working memory tasks with domain-specific distractor trials inserted into each interstimulus interval of the n-back tasks. In two experiments, object n-back performance demonstrated greater sensitivity to verbal distraction, whereas spatial n-back performance demonstrated greater sensitivity to motion distraction. Visual object and spatial working memory may differ fundamentally in that the mnemonic representation of featural characteristics of objects incorporates a verbal (perhaps semantic) code, whereas the mnemonic representation of the location of objects does not. Thus, the processes supporting working memory for these two types of information may differ in more ways than those dictated by the "what/where" organization of the visual system, a fact more easily reconciled with a component process than a memory systems account of working memory function.     

论执行功能及其发展研究

执行功能(executive function,EF)是一个还没有明确定义的概念,但又是一个重要的概念,通常是指个体对思想和行动进行有意识控制的心理过程。目前的主要观点包括神经心理学观点、抑制控制理论、高级认知能力理论(包括工作记忆理论、抑制及工作记忆理论)和认知复杂性和控制理论。主要的研究任务类型包括卡片分类任务、延迟反应任务、基于Stroop任务的冲突反应模式任务、心理理论任务、问题解决任务、“热”执行功能任务等。未来的研究应着重弄清执行功能的本质,结合认知神经科学阐明其与脑神经结构和功能的关系,并从区分“冷”执行功能和“热”执行功能的角度,探讨以情感激活为特点的“热认知”在执行功能中的作用。     

The role of memory representation in the vigilance decrement

Working memory load is critically important for the overall level of performance on vigilance tasks. However, its role in a key aspect of vigilance—sensitivity decrement over time—is unclear. We used a dual-task procedure in which either a spatial or a nonspatial working memory task was performed simultaneously with a spatial vigilance task for 20 min. Sensitivity in the vigilance task declined over time when the concurrent task involved spatial working memory. In contrast, there was no sensitivity decrement with a nonspatial working memory task. The results provide the first evidence of a specific role for working memory representation in vigilance decrement. The findings are also consistent with a multiple resource theory in which separate resources for memory representation and cognitive control operations are differentially susceptible to depletion over time, depending on the demands of the task at hand.     

Spatial memory in rats after 25 hours

We investigated the time course of spatial-memory decay in rats using an eight-arm radial maze. It is well established that performance remains high with retention intervals as long as 4 h, but declines to chance with a 24-h retention interval (Beatty, W. W., & Shavalia, D. A. (1980b). Spatial memory in rats: time course of working memory and effect of anesthetics. Behavioral & Neural Biology, 28, 454-462). It is possible that 24 h reflects a genuine retention limitation of rat spatial memory. Alternatively, it may be possible to identify factors that might support memory performance even after very long delays. The current experiment was conducted to test the above two hypotheses. We evaluated performance using two intertrial intervals (24 and 48 h) and two retention intervals (1 and 25 h). Increasing the intertrial interval produced an approximately constant increase in performance for both retention intervals. This improvement is consistent with a trial-spacing effect (i.e., the superiority of spaced over massed trials). Rat spatial memory apparently lasts at least 25 h.     

Allocentric spatial learning by hippocampectomised rats: A further test of the “spatial mapping” and “working memory” theories of hippocampal function

This paper reports a series of three experiments that tested the “spatial-mapping” and “working-memory” theories of hippocampal function. The experimental designs incorporate separate reference- and working-memory procedures of a water-escape task, using both spatial and non-spatial learning. In Experiment 1 (Reference memory), rats with hippocampal (HC) or cortical (CC) lesions and unoperated (UNOP) rats learned to swim to a rigid visible escape platform while avoiding contact with a floating one. In the nonspatial task, the platforms each occupied any of 8 possible positions in the pool over successive trials but differed in appearance. In the spatial task, the platforms were of identical appearance but the safe one always occupied a single fixed location. The HC rats showed a highly specific spatial learning impairment but did learn to perform consistently above chance towards the end of training. In Experiment 2 (working memory), new groups of rats were trained on similar spatial and nonspatial tasks, but the platform designated correct-in terms of its visual appearance or its spatial location-was randomly changed each day. No animal learned the nonspatial task despite extensive training. Performance on the spatial version unexpectedly revealed an impairment in the CC as well as the HC group relative to the UNOP rats. However, the HCs again performed at above chance levels and demonstrated rapid (I-trial) spatial learning towards the end training. Experiment 3 used a place navigation matching-to-sample task examine spatial working memory further. Each day, an underwater platform was hidden at any of 4 possible locations, and the rats were given 2 trials to search for it. Both UNOP and CC rats located the platform faster on Trial 2 than on Trial 1, even when the inter-trial interval was long as 30min. HC rats were no faster on Trial 2 than on Trial 1. We conclude that hippocampal lesions (1) severely but partially impair spatial but not visual reference memory and (2) give rise to different patterns impairment in different working-mermory tasks. The results are a chal lenge to both the spatial-mapping and working-memory theories.     

Cognitive decline and human (Homo sapiens) aging: an investigation using a comparative neuropsychological approach

Using a comparative neuropsychological approach, the authors compared performance of younger and healthy older adults ages 65 and over on tasks originally developed to measure cognition in animals. A battery of 6 tasks was used to evaluate object discrimination, egocentric spatial abilities, visual and spatial working memory, and response shifting. Older adults performed more poorly than younger adults on tasks that evaluate egocentric spatial abilities, response shifting, and to a lesser extent object recognition. The two groups did not differ for tasks that evaluate spatial working memory and object discrimination. The impairments the authors observed in tasks that evaluate response shifting and object recognition are consistent with those found in canines and primates as well as those found in Alzheimer's disease. The results are consistent with the notion that cognitive processes supported by the amygdala and the orbitofrontal cortex are among the first to decline with increasing age in both humans and animals.     

A male advantage for spatial and object but not verbal working memory using the n-back task

Sex-related differences have been reported for performance and neural substrates on some working memory measures that carry a high cognitive load, including the popular n-back neuroimaging paradigm. Despite some evidence of a sex effect on the task, the influence of sex on performance represents a potential confound in neuroimaging research. The present study investigated sex-related differences in verbal, spatial, and common object versions of the high cognitive load "n-back" working memory task. Eighteen male and 18 female undergraduates completed all 3 versions of the task. A mixed ANOVA, with Sex (male and female) as the between-subjects factor and Condition (verbal, spatial, and object) as the within-subjects repeated measure revealed that males were significantly more accurate than females on the spatial and object versions of the n-back task and performed equivalently to females on the verbal version of the task. Although the expected female advantage for verbal working memory was not found using this effortful n-back task, these results support a male advantage for high cognitive load spatial and object working memory. Future research should take into account the influence of sex on performance of the n-back task, and examine sex-related differences in working memory using other paradigms.     

Young adults’ reactions to infant crying

AimTo determine whether young childless adults show negative emotions and cognitive disturbances when listening to infant crying, compared to other disturbing noises, and whether negative emotions and cognitive disturbances are associated.MethodsWe tested the cognitive performances and emotional reactions of 120 childless participants on a working memory task while being subjected to different disturbing noises including infant crying.ResultsParticipants had the least correct trials on the working memory task, and showed the most negative emotions, when hearing infant crying as compared to the other noises. Participants also showed less positive emotions when hearing infant crying as compared to working in silence. Overall, negative emotions were associated with less correct trials on the working memory task, except in the infant crying condition. Furthermore, cognitive performance and emotional reactions to infant crying were unrelated to personality characteristics.ConclusionNegative emotions and cognitive disturbances may be general adult responses to infant crying that are not limited to parents. These results suggest a broadly present human emotional and cognitive response to infant crying, that may underlie a general predisposition to care for infants in distress.     

Olfactory working memory: effects of verbalization on the 2-back task

Working memory for odors, which has received almost no attention in the literature, was investigated in two experiments. We show that performance in a 2-back task with odor stimuli is well above chance. This is true not only for highly familiar odors, as has been shown by Dade, Zatorre, Evans, and Jones-Gotman, NeuroImage, 14, 650–660, (2001), but also for unfamiliar ones that are notoriously difficult to name. We can conclude that information about an olfactory stimulus can be retained in the short term and can continuously be updated for comparison with new olfactory probes along the lines of a functional odor working memory. However, the performance in the working memory task is highly dependent on participants’ verbalization of the odor. In addition, results indicated that odor working memory performance is dependent on the ability to discriminate among the odor stimuli (Experiment 2). The results are discussed in relation to recent ideas of a separate olfactory working memory slave system.     

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.     

Oculomotor delayed response abnormalities in young offspring and siblings at risk for schizophrenia

Individuals with schizophrenia are know to demonstrate cognitive and behavioral difficulties, particularly alterations in executive functions, including working memory. It is unclear whether these deficits reflect trait-related vulnerability to schizophrenia indicators and can be assessed by studying nonpsychotic relatives of patients with schizophrenia. In this study, we used an oculomotor delayed response (ODR) paradigm to examine spatial working memory in 37 "high-risk" child and adolescent offspring and siblings (age range=6-25 years) of patients with schizophrenia or schizoaffective disorder. Compared with 37 age- and sex-matched healthy controls (age range=6-23 years), high-risk subjects showed nonsignificantly greater errors in the ODR task at longer delay intervals. Statistical analyses suggested that performance improved with age in healthy control subjects, whereas it worsened with age in high-risk subjects. In both groups, the ODR errors were generally associated with poorer sustained attention (Continuous Performance Test: visuospatial d prime), somewhat poorer executive function (Wisconsin Card Sorting Test), and elevated Heinrichs-Buchanan neurological soft signs scores. These findings indicate the presence of spatial working memory abnormalities in individuals at risk for schizophrenia. Furthermore, these abnormalities may be progressive in nature. The ODR task is a valuable indicator of prefrontal cortical function and spatial working memory and may be a potentially valuable marker for familial risk of schizophrenia.     

A model of working memory capacity in the radial-arm maze task

The radial arm maze is one of the most commonly used tests for assessing working memory in laboratory animals. However, to date, there exists no quantitative method of estimating working memory capacity from performance on this task. Here, we present a mathematical model of performance on the radial arm maze from which we can derive estimates of capacity. We derive explicit results for the two most commonly used measures of performance as functions of number of arms in the maze and memory capacity, assuming a uniform random search. We simulate random non-uniform search strategies. Comparing our model to previous experiments, we show that our model predicts a working memory capacity in the range of 3-9 at the level of performance observed in these experiments. This estimate is within the typical estimate of human working memory capacity. Performance of rats on large mazes (e.g. 48 arms) has been used as evidence that the working memory capacity of rats may be significantly larger than that of humans. We report that memory capacity in the range 3-9 is sufficient to explain the performance of rats in very large radial mazes. Furthermore, when we simulate non-uniform random search strategies observed in the experiments, the resulting estimates do not differ significantly from those assuming a uniform random search. We conclude that a list-based model of working memory with modest capacity is more powerful than previously expected.     

行动控制风格对情绪与空间工作记忆关系的调节效应

以90名大学生为被试,探讨行动控制风格对情绪与空间工作记忆关系的影响。采用3(情绪体验:中性、积极、消极)×2(行动控制风格:行动导向、状态导向)混合实验设计,进行延迟匹配记忆任务实验。研究发现,中性与积极情绪体验下,行动导向与状态导向空间工作记忆的反应时与正确率均无显著差异; 消极情绪体验下,行动导向与状态导向空间工作记忆结果差异显著,行动导向的反应时更短,正确率更高。结果表明,消极情绪体验下,行动控制风格对情绪与空间工作记忆的关系起调节作用。     

Constructing visual representations of natural scenes: the roles of short- and long-term visual memory

A "follow-the-dot" method was used to investigate the visual memory systems supporting accumulation of object information in natural scenes. Participants fixated a series of objects in each scene, following a dot cue from object to object. Memory for the visual form of a target object was then tested. Object memory was consistently superior for the two most recently fixated objects, a recency advantage indicating a visual short-term memory component to scene representation. In addition, objects examined earlier were remembered at rates well above chance, with no evidence of further forgetting when 10 objects intervened between target examination and test and only modest forgetting with 402 intervening objects. This robust prerecency performance indicates a visual long-term memory component to scene representation.     

A facilitative effect of negative affective valence on working memory

Gotoh, F., Kikuchi, T. &Olofsson, U. (2010). A facilitative effect of negative affective valence on working memory. Scandinavian Journal of Psychology, 51, 185–191. Previous studies have shown that negatively valenced information impaired working memory performance due to an attention‐capturing effect. The present study examined whether negative valence could also facilitate working memory. Affective words (negative, neutral, positive) were used as retro‐cues in a working memory task that required participants to remember colors at different spatial locations on a computer screen. Following the cue, a target detection task was used to either shift attention to a different location or keep attention at the same location as the retro‐cue. Finally, participants were required to discriminate the cued color from a set of distractors. It was found that negative cues yielded shorter response times (RTs) in the attention‐shift condition and longer RTs in the attention‐stay condition, compared with neutral and positive cues. The results suggest that negative affective valence may enhance working memory performance (RTs), provided that attention can be disengaged.     

Performance dissociation during verbal and spatial working memory tasks

Past research has inconsistently distinguished the neural substrates of various types of working memory. Task design and individual performance differences are known to alter patterns of brain response during working-memory tasks. These task and individual differences may have produced discrepancies in imaging findings. This study of 50 healthy adults (M(age) = 19.6 yr., SD = .8) examined performance during various parametric manipulations of a verbal and spatial n-back working-memory task. Performance systematically dissociated on the basis of working-memory load, working memory type, and stimulus difficulty, with participants having greater accuracy but slower response time during conditions requiring verbal versus spatial working memory. These findings hold implications for cognitive and neuroimaging studies of verbal and spatial working memory and highlight the importance of considering both task design and individual behavior.     

Susceptibility of spatial and verbal working memory to demands of the central executive

We used a dual-task paradigm to examine the degree to which domain-specific spatial and verbal subsystems depend on the domain-general central executive. Forty participants were asked to retain spatial or verbal information while performing a concurrent secondary task related to simple arithmetic. The secondary tasks consisted of three cognitive processes: single-digit addition, a digit-carrying operation, and digit reading. The single-digit addition and carry operation include central executive functioning, while digit reading relies solely on the phonological loop. The single-digit addition caused a performance decrement on the spatial working memory task, while the digit reading impaired performance on the verbal working memory task. The carry operation interfered with recall accuracy on both working memory tasks. The spatial working memory task was significantly correlated with the verbal working memory task only when the secondary task was more demanding on the central executive. Our results suggest that spatial working memory rather than verbal working memory is susceptible to failure of central executive functioning and that the central executive plays an important role in regulating the cognitive demands of different domains.