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.     

A Sex Difference on a Novel Spatial Working Memory Task in Humans

Neurophysiological and behavioral evidence suggests that the prefrontal cortex (PFC) may be sexually differentiated in nonhuman primates. The present study examined whether there are sex differences in working memory in humans that might reflect sexual differentiation of human PFC. Male and female undergraduates were administered a novel multitrial spatial working memory task (SPWM) and a verbal working memory task. In three experiments, females committed significantly fewer working memory errors and took significantly less time to reach criterion than males on the SPWM task. The female advantage was not accounted for by differences in general intellectual ability, attention, perceptual speed, incidental memory, or speed of verbal access. In Study 3, a sex difference was also observed on a measure of verbal working memory. The findings suggest that some prefrontal functions may be sexually differentiated in humans.     

Sex differences in object location memory and spatial navigation in Long-Evans rats

In both humans and rodents, males typically excel on a number of tasks requiring spatial ability. However, human females exhibit advantages in memory for the spatial location of objects. This study investigated whether rats would exhibit similar sex differences on a task of object location memory (OLM) and on the watermaze (WM). We predicted that females should outperform males on the OLM task and that males should outperform females on the WM. To control for possible effects of housing environment, rats were housed in either complex environments or in standard shoebox housing. Eighty Long-Evans rats (40 males and 40 females) were housed in either complex (Complex rats) or standard shoebox housing (Control rats). Results indicated that males had superior performance on the WM, whereas females outperformed males on the OLM task, regardless of housing environment. As these sex differences cannot be easily attributed to differences in cognitive style related to linguistic processing of environmental features or to selection pressures related to the hunting gathering evolutionary prehistory of humans, these data suggest that sex differences in spatial ability may be related to traits selected for by polygynous mating strategies.     

Book Reviews

There is long-standing evidence for verbal working memory impairments in both children and adults with dyslexia. By contrast, spatial memory appears largely to be unimpaired. In an attempt to distinguish between phonological and central executive accounts of the impairments in working memory, a set of phonological and spatial working memory tasks was designed to investigate the key issues in working memory, task type, task demands (static, dynamic, and updating), and task complexity. Significant differences emerged between the dyslexic and nondyslexic participants on the verbal working memory tasks employed in Experiment 1, thereby providing further evidence for continuing dyslexic impairments of working memory into adulthood. The nature of the deficits suggested a problem with the phonological loop, with there being little evidence to implicate an impairment of the central executive. Due to the difficulties associated with separating verbal working memory and phonological processing, however, performance was investigated in Experiment 2 using visuospatial measures of working memory. The results of the visuospatial tasks indicated no between-group differences in static spatial memory, which requires the short-term storage of simultaneously presented information. In almost all conditions there were no between-group differences in dynamic spatial memory that demands the recall of both location and order of stimuli presented sequentially. However, a significant impairment occurred on the dynamic task under high memory updating load, on which dyslexic adults showed nonphonological working memory deficits. In the absence of an explanation involving verbal recoding, this finding is interpreted in terms of a central executive or automaticity impairment in dyslexia.     

GENDER AND TASK IN THE DETERMINATION OF SPATIAL COGNITIVE PERFORMANCE

A variety of sex differences in spatial cognitive performance have been documented. However, factors other than those specifically related to gender and cognition per se, such as the perceived spatial character of given tasks, may contribute to such differences. In the present experiments, spatial memory and mental image rotation tasks were presented to female and male adults. The task formats or instructions were varied to emphasize or deemphasize the spatial character of the tasks. Highly "spatial" instructions or format significantly depressed performance on spatial tasks for women but not for men. "Nonspatial" instructions or format, within which the spatial character of the task was not explicit, resulted in no significant differences between the performances of women and men on either type of task. These findings indicate that instructional or format effects relating to the purported "spatial" character of a given task may significantly influence the relative performance of women and men.     

Sex differences in the processing of flankers

The study of sex differences in cognition has often focused on differences in spatial processing. Recently, sex differences in selective attention have been observed by Bayliss, di Pellegrino, and Tipper (2005), showing that women are more influenced than men by irrelevant spatial cues. The current study elaborates on this finding and tests whether sex differences in the processing of irrelevant information also occur in a simpler task, in which there is no need to redirect visual attention and no need to remember multiple spatial stimulus–response associations. Here, attention is studied using a novel combination of a go/no-go task and a flanker task. A total of 80 neurotypical participants were studied, and it was found that responses in women were more strongly affected by flanker information than were responses in men. This suggests that these sex differences were not due to difficulties with spatial reorientation, or remembering spatial stimulus–response relationships. The findings are discussed in the context of the hunter–gatherer theory of sex differences.     

Cultural differences in visual attention: Implications for distraction processing

We investigated differences between participants of East Asian and Western descent in attention to and implicit memory for irrelevant words which participants were instructed to ignore while completing a target task (a Stroop Task in Experiment 1 and a 1‐back task on pictures in Experiment 2 ). Implicit memory was measured using two conceptual priming tasks (category generation in Experiment 1 and general knowledge in Experiment 2 ). Participants of East Asian descent showed reliable implicit memory for previous distractors relative to those of Western descent with no evidence of differences on target task performance. We also found differences in a Corsi Block spatial memory task in both studies, with superior performance by the East Asian group. Our findings suggest that cultural differences in attention extend to task‐irrelevant background information, and demonstrate for the first time that such information can boost performance when it becomes relevant on a subsequent task.     

Sex differences in cognition among illiterate Bangladeshis: a comparison with literate Bangladeshis and Swedes

We investigated the presence and magnitude of sex differences in late adulthood, assessing 426 illiterate Bangladeshis, 239 literate Bangladeshis, and 598 Swedes. The cognitive domains examined included calculation, episodic memory, spatial visualization, and global cognitive ability. In general, men performed at a higher level than women on tasks assessing calculation and spatial visualization, whereas women performed at a higher level than men on the episodic memory task. Notably, the pattern of cognitive sex differences was similar irrespective of nationality and literacy, although the magnitude of the male advantage was inversely related to level of education. Finally, the low performance of the illiterate women demonstrated the penalizing effect restrictions in public exposure might have on cognitive performance.     

Sex differences in a human analogue of the Radial Arm Maze: the "17-Box Maze Test"

This study investigated sex differences in spatial memory using a human analogue of the Radial Arm Maze: a revision on the Nine Box Maze originally developed by called the 17-Box Maze Test herein. The task encourages allocentric spatial processing, dissociates object from spatial memory, and incorporates a within-participants design to provide measures of location and object, working and reference memory. Healthy adult males and females (26 per group) were administered the 17-Box Maze Test, as well as mental rotation and a verbal IQ test. Females made significantly fewer errors on this task than males. However, post hoc analysis revealed that the significant sex difference was specific to object, rather than location, memory measures. These were medium to large effect sizes. The findings raise the issue of task- and component-specific sexual dimorphism in cognitive mapping.     

Differentiating between verbal and spatial encoding using eye-movement recordings

Visual information processing is guided by an active mechanism generating saccadic eye movements to salient stimuli. Here we investigate the specific contribution of saccades to memory encoding of verbal and spatial properties in a serial recall task. In the first experiment, participants moved their eyes freely without specific instruction. We demonstrate the existence of qualitative differences in eye-movement strategies during verbal and spatial memory encoding. While verbal memory encoding was characterized by shifting the gaze to the to-be-encoded stimuli, saccadic activity was suppressed during spatial encoding. In the second experiment, participants were required to suppress saccades by fixating centrally during encoding or to make precise saccades onto the memory items. Active suppression of saccades had no effect on memory performance, but tracking the upcoming stimuli decreased memory performance dramatically in both tasks, indicating a resource bottleneck between display-controlled saccadic control and memory encoding. We conclude that optimized encoding strategies for verbal and spatial features are underlying memory performance in serial recall, but such strategies work on an involuntary level only and do not support memory encoding when they are explicitly required by the task.     

No sex differences in spatial location memory for abstract designs

Previous research has demonstrated a female advantage, albeit imperfectly, on tests of object location memory where object identity information is readily available. However, spatial and visual elements are often confounded in the experimental tasks used. Here spatial and visual memory performance was compared in 30 men and 30 women by presenting 12 abstract designs in a spatial array for recall and recognition (visual memory) and spatial location ("object" location memory). Object location memory was measured via a sensitive absolute displacement score defined as the distance in mms between the position assigned to the object during recall and the actual position it originally occupied. There were no sex differences in either the visual or spatial location tests. Controlling for age and estimated IQ scores made no impact on the results. These data suggest an absence of a sex difference in purely visual and spatial aspects of object location memory.     

Sex differences in visuospatial working memory: components of cognitive processing

Numerous studies have shown that sex differences in visuospatial tasks vary in size and direction depending on the nature of the task, with large differences favoring males on tasks that require transformations in visuospatial working memory. The cognitive processes underlying these differences were investigated using laboratory tasks developed by Dror and Kosslyn (1994). Four cognitive components of visuospatial working memory were assessed—image generation, maintenance, scanning, and transformation—in an attempt to identify the components that would show differential effects for females and males. The image generation task required retrieval of shape information from long-term memory, generation of a visual image in working memory, and utilization of the information about the shape in a decision task. The image maintenance task required only the latter two processes. The information processing demands required by scanning and rotation tasks came from the need to transform the visual image so that it could be used in decision making. Males responded more quickly on all four tasks (ds between .63 and .77), with no between-sex differences in accuracy. We concluded that speed of processing is central to understanding sex differences in visuospatial working memory. We discuss implications of these findings for performance on real-world visuospatial tasks.     

Sex differences in visual-spatial working memory: A meta-analysis

Visual-spatial working memory measures are widely used in clinical and experimental settings. Furthermore, it has been argued that the male advantage in spatial abilities can be explained by a sex difference in visual-spatial working memory. Therefore, sex differences in visual-spatial working memory have important implication for research, theory, and practice, but they have yet to be quantified. The present meta-analysis quantified the magnitude of sex differences in visual-spatial working memory and examined variables that might moderate them. The analysis used a set of 180 effect sizes from healthy males and females drawn from 98 samples ranging in mean age from 3 to 86 years. Multilevel meta-analysis was used on the overall data set to account for non-independent effect sizes. The data also were analyzed in separate task subgroups by means of multilevel and mixed-effects models. Results showed a small but significant male advantage (mean d = 0.155, 95 % confidence interval = 0.087-0.223). All the tasks produced a male advantage, except for memory for location, where a female advantage emerged. Age of the participants was a significant moderator, indicating that sex differences in visual-spatial working memory appeared first in the 13-17 years age group. Removing memory for location tasks from the sample affected the pattern of significant moderators. The present results indicate a male advantage in visual-spatial working memory, although age and specific task modulate the magnitude and direction of the effects. Implications for clinical applications, cognitive model building, and experimental research are discussed.     

An fMRI study of sex differences in regional activation to a verbal and a spatial task

Sex differences in cognitive performance have been documented, women performing better on some phonological tasks and men on spatial tasks. An earlier fMRI study suggested sex differences in distributed brain activation during phonological processing, with bilateral activation seen in women while men showed primarily left-lateralized activation. This blood oxygen level-dependent fMRI study examined sex differences (14 men, 13 women) in activation for a spatial task (judgment of line orientation) compared to a verbal-reasoning task (analogies) that does not typically show sex differences. Task difficulty was manipulated. Hypothesized ROI-based analysis documented the expected left-lateralized changes for the verbal task in the inferior parietal and planum temporal regions in both men and women, but only men showed right-lateralized increase for the spatial task in these regions. Image-based analysis revealed a distributed network of cortical regions activated by the tasks, which consisted of the lateral frontal, medial frontal, mid-temporal, occipitoparietal, and occipital regions. The activation was more left lateralized for the verbal and more right for the spatial tasks, but men also showed some left activation for the spatial task, which was not seen in women. Increased task difficulty produced more distributed activation for the verbal and more circumscribed activation for the spatial task. The results suggest that failure to activate the appropriate hemisphere in regions directly involved in task performance may explain certain sex differences in performance. They also extend, for a spatial task, the principle that bilateral activation in a distributed cognitive system underlies sex differences in performance.     

Long-term memory of hierarchical relationships in free-living greylag geese

Animals may memorise spatial and social information for many months and even years. Here, we investigated long-term memory of hierarchically ordered relationships, where the position of a reward depended on the relationship of a stimulus relative to other stimuli in the hierarchy. Seventeen greylag geese (Anser anser) had been trained on discriminations between successive pairs of five or seven implicitly ordered colours, where the higher ranking colour in each pair was rewarded. Geese were re-tested on the task 2, 6 and 12 months after learning the dyadic colour relationships. They chose the correct colour above chance at all three points in time, whereby performance was better in colour pairs at the beginning or end of the colour series. Nonetheless, they also performed above chance on internal colour pairs, which is indicative of long-term memory for quantitative differences in associative strength and/or for relational information. There were no indications for a decline in performance over time, indicating that geese may remember dyadic relationships for at least 6 months and probably well over 1 year. Furthermore, performance in the memory task was unrelated to the individuals’ sex and their performance while initially learning the dyadic colour relationships. We discuss possible functions of this long-term memory in the social domain.     

Visual working memory as the substrate for mental rotation

In mental rotation, a mental representation of an object must be rotated while the actual object remains visible. Where is this representation stored while it is being rotated? To answer this question, observers were asked to perform a mental rotation task during the delay interval of a visual working memory task. When the working memory task required the storage of object features, substantial bidirectional interference was observed between the memory and rotation tasks, and the interference increased with the degree of rotation. However, rotation-dependent interference was not observed when a spatial working memory task was used instead of an object working memory task. Thus, the object working memory subsystem—not the spatial working memory subsystem—provides the buffer in which object representations are stored while they undergo mental rotation. More broadly, the nature of the information being stored—not the nature of the operations performed on this information—may determine which subsystem stores the information.     

Effects of Domain,Retention Interval,and Information Load on Young and Older Adults’ Visuospatial Working Memory

Age-related differences in visuospatial working memory were examined in 69 young adults and 49 older adults exposed to three pairs of tasks. Each pair consisted of one task involving information about the form or appearance of items and another task involving information about item locations. The first pair of tasks manipulated retention interval and required maintaining information about one item. The second pair also manipulated retention interval and required maintaining information about multiple items presented simultaneously. The third pair manipulated the number of sequentially presented items. Analyses of the first two pairs of tasks revealed significant age deficits in working memory for spatial locations but not in working memory for visual features. Notably, there were no age differences in the effect of retention interval on any of the four tasks, suggesting that visuospatial information is lost at similar rates in older and young adults. Analyses of the third pair of tasks revealed that, regardless of domain, increasing the amount of information impaired older adults’ memory performance to a greater extent than young adults’ performance. Thus, the present results suggest differences in basic working memory capacity in both domains, but a lack of age differences in rates of forgetting from working memory, and greater age-related deficits in the spatial domain than in the object domain.     

Preserved spatial memory over brief intervals in older adults

Two studies compared young and older adults' memory for location information after brief intervals. Experiment 1 found that accuracy of intentional spatial memory for individual locations was similar in young and older participants for set sizes of 3 and 6. Both groups also encoded individual locations in relation to the larger configuration of locations. Experiment 2 showed that like young adults, older adults' latency to respond to a test probe in a letter working memory task was negatively influenced by spatial information that was irrelevant to the task. This interference effect indicated preserved incidental memory for spatial information in older adults. Together, these data suggest that initial encoding of spatial information for relatively small numbers of items is largely preserved in healthy older adults and that representations of spatial information persist over short intervals.     

Visual search is slowed when visuospatial working memory is occupied

Visual working memory plays a central role in most models of visual search. However, a recent study showed that search efficiency was not impaired when working memory was filled to capacity by a concurrent object memory task (Woodman, Vogel, & Luck, 2001). Objects and locations may be stored in separate working memory subsystems, and it is plausible that visual search relies on the spatial subsystem, but not on the object subsystem. In the present study, we sought to determine whether maintaining spatial information in visual working memory impairs the efficiency of a concurrent visual search task. Visual search efficiency and spatial memory accuracy were both impaired when the search and the memory tasks were performed concurrently, as compared with when the tasks were performed separately. These findings suggest that common mechanisms are used to process information during difficult visual search tasks and to maintain spatial information in working memory.     

Central executive involvement in children's spatial memory

Previous research with adults found that spatial short-term and working memory tasks impose similar demands on executive resources. We administered spatial short-term and working memory tasks to 8- and 11-year-olds in three separate experiments. In Experiments 1 and 2 an executive suppression task (random number generation) was found to impair performances on a short-term memory task (Corsi blocks), a working memory task (letter rotation), and a spatial visualisation task (paper folding). In Experiment 3 an articulatory suppression task only impaired performance on the working memory task. These results suggest that short-term and working memory performances are dependent on executive resources. The degree to which the short-term memory task was dependent on executive resources was expected to be related to the amount of experience children have had with such tasks. Yet we found no significant age-related suppression effects. This was attributed to differences in employment of cognitive strategies by the older children.