How rats perform spatial working memory tasks: limitations in the use of egocentric and idiothetic working memory

Rats of the Dark Agouti strain were trained on delayed alternation under conditions that should encourage egocentric working memory. In two experiments a T-maze was set within a cross-maze so that different arms could be used for the sample and test runs. The maze had high opaque side-walls, and testing was conducted in low light levels so that distal visual cues might be eliminated. By rotating the maze 90° between the sample and choice run and by using two identical mazes set side by side it was possible to nullify other spatial strategies. Experiments 1 and 2 showed that rats preferentially used place information, intramaze cues, and direction cues, even though only egocentric or idiothetic (nonmatch-to-turn) working memory could successfully solve every trial. Rats were able to maintain an accurate sense of location within the maze even though distal cues were not visible and the animal was moved between the sample and choice runs. Experiment 2 confirmed that another rat strain (Long-Evans) shows the same learning profiles. Both experiments indicate that rats are very poor at using either egocentric or idiothetic information to alternate, and that retention delays as short as 10 s can eliminate the use of these forms of memory.     

How rats perform spatial working memory tasks: Limitations in the use of egocentric and idiothetic working memory

Rats of the Dark Agouti strain were trained on delayed alternation under conditions that should encourage egocentric working memory. In two experiments a T-maze was set within a cross-maze so that different arms could be used for the sample and test runs. The maze had high opaque side-walls, and testing was conducted in low light levels so that distal visual cues might be eliminated. By rotating the maze 90° between the sample and choice run and by using two identical mazes set side by side it was possible to nullify other spatial strategies. Experiments 1 and 2 showed that rats preferentially used place information, intramaze cues, and direction cues, even though only egocentric or idiothetic (nonmatch-to-turn) working memory could successfully solve every trial. Rats were able to maintain an accurate sense of location within the maze even though distal cues were not visible and the animal was moved between the sample and choice runs. Experiment 2 confirmed that another rat strain (Long-Evans) shows the same learning profiles. Both experiments indicate that rats are very poor at using either egocentric or idiothetic information to alternate, and that retention delays as short as 10 s can eliminate the use of these forms of memory.     

Executive processes in visual and spatial working memory tasks

Three experiments are reported, which have investigated the nature of the cognitive mechanisms that underlie performance on specific visuo-spatial working memory tasks, with the emphasis on exploring the extent of central executive involvement. Experiments 1 and 2 employed oral random digit generation as an executive task within a dual-task paradigm. The results of both experiments indicated that visuo-spatial tasks that involve sequential processing of information show more interference with random digit generation than do visuo-spatial tasks that involve simultaneous processing. The third experiment substituted oral random digit generation for executive tasks that did not involve memory for serial order (vigilance tasks adapted from Vandierendonck, De Vooght, & Van der Goten, 1998b). The results indicated significant interference between the vigilance tasks and the sequential visuo-spatial task, but not with the simultaneous visuo-spatial task. Overall the results of the three experiments are interpreted as indicating that serial sequential visuo-spatial tasks involve executive resources to a significantly greater extent than do simultaneous visuo-spatial tasks, and that this can have implications for studies that attempt to make use of such tasks to fractionate separable visual and spatial components within working memory.     

Executive processes in visual and spatial working memory tasks

Three experiments are reported, which have investigated the nature of the cognitive mechanisms that underlie performance on specific visuo-spatial working memory tasks, with the emphasis on exploring the extent of central executive involvement. Experiments 1 and 2 employed oral random digit generation as an executive task within a dual-task paradigm. The results of both experiments indicated that visuo-spatial tasks that involve sequential processing of information show more interference with random digit generation than do visuo-spatial tasks that involve simultaneous processing. The third experiment substituted oral random digit generation for executive tasks that did not involve memory for serial order (vigilance tasks adapted from Vandierendonck, De Vooght, & Van der Goten, 1998b). The results indicated significant interference between the vigilance tasks and the sequential visuo-spatial task, but not with the simultaneous visuo-spatial task. Overall the results of the three experiments are interpreted as indicating that serial sequential visuo-spatial tasks involve executive resources to a significantly greater extent than do simultaneous visuo-spatial tasks, and that this can have implications for studies that attempt to make use of such tasks to fractionate separable visual and spatial components within working memory.     

Language, space, and the development of cognitive flexibility in humans: the case of two spatial memory tasks

Prior experiments have shown that young children, like adult rats, rely mainly on information about the macroscopic shape of the environment to reorient themselves, whereas human adults rely more flexibly on combinations of spatial and non-spatial landmark information. Adult rats have also been shown to exhibit a striking limitation in another spatial memory task, movable object search, again a limitation not shown by human adults. The present experiments explored the developmental change in humans leading to more flexible, human adult-like performance on these two tasks. Experiment 1 identified the age range of 5-7 years as the time the developmental change for reorientation occurs. Experiment 2 employed a multiple regression approach to determine that among several candidate measures, only a specific language production measure, the production of phrases specifying exactly the information needed to solve the task like adults, correlated with the reorientation performance of children in this age range. Experiment 3 revealed that similar language production abilities were associated with more flexible moving object search task performance. These results, in combination with findings with human adults, suggest that language production skills play a causal role in allowing older humans to construct novel representations rapidly, which can then be used to transcend the limits of phylogenetically older cognitive processes.     

Verbal cues flexibly transform spatial representations in human memory

ABSTRACT

Humans possess a unique ability to communicate spatially-relevant information, yet the intersection between language and navigation remains largely unexplored. One possibility is that verbal cues accentuate heuristics useful for coding spatial layouts, yet this idea remains largely untested. We test the idea that verbal cues flexibly accentuate the coding of heuristics to remember spatial layouts via spatial boundaries or landmarks. The alternative hypothesis instead conceives of encoding during navigation as a step-wise process involving binding lower-level features, and thus subsequently formed spatial representations should not be modified by verbal cues. Across three experiments, we found that verbal cues significantly affected pointing error patterns at axes that were aligned with the verbally cued heuristic, suggesting that verbal cues influenced the heuristics employed to remember object positions. Further analyses suggested evidence for a hybrid model, in which boundaries were encoded more obligatorily than landmarks, but both were accessed flexibly with verbal instruction. These findings could not be accounted for by a tendency to spend more time facing the instructed component during navigation, ruling out an attentional-encoding mechanism. Our findings argue that verbal cues influence the heuristics employed to code environments, suggesting a mechanism for how humans use language to communicate navigationally-relevant information.     

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.     

Bonobos and orangutans,but not chimpanzees,flexibly plan for the future in a token-exchange task

Non-human animals, including great apes, have been suggested to share some of the skills for planning that humans commonly exhibit. A crucial difference between human and non-human planning may relate to the diversity of domains and needs in which this skill is expressed. Although great apes can save tools for future use, there is little evidence yet that they can also do so in other contexts. To investigate this question further, we presented the apes with a planning token-exchange task that differed from standard tool-use tasks. Additionally, we manipulated the future outcome of the task to investigate planning flexibility. In the Exchange condition, subjects had to collect, save and transport tokens because they would need them 30 min later to exchange them for food with a human, i.e., “bring-back” response. In the Release condition, the collection and transport of tokens were not needed as no exchange took place after 30 min. Out of 13 subjects, eight solved the task at least once in the Exchange condition, with chimpanzees appearing less successful than the other species. Importantly, three individuals showed a clear differential response between conditions by producing more “bring-back” responses in the Exchange than in the Release conditions. Those bonobo and orangutan individuals hence adapted their planning behavior according to changing needs (i.e., they brought tokens back significantly more often when they would need them). Bonobos and orangutans, unlike chimpanzees, planned outside the context of tool-use, thus challenging the idea that planning in these species is purely domain-specific.     

The use of prosodic cues in language discrimination tasks by rats

Recent research with cotton-top tamarin monkeys has revealed language discrimination abilities similar to those found in human infants, demonstrating that these perceptual abilities are not unique to humans but are also present in non-human primates. Specifically, tamarins could discriminate forward but not backward sentences of Dutch from Japanese, using both natural and synthesized utterances. The present study was designed as a conceptual replication of the work on tamarins. Results show that rats trained in a discrimination learning task readily discriminate forward, but not backward sentences of Dutch from Japanese; the results are particularly robust for synthetic utterances, a pattern that shows greater parallels with newborns than with tamarins. Our results extend the claims made in the research with tamarins that the capacity to discriminate languages from different rhythmic classes depends on general perceptual abilities that evolved at least as far back as the rodents. Electronic Publication     

Geometric cues,reference frames,and the equivalence of experienced-aligned and novel-aligned views in human spatial memory

Spatial memories are often organized around reference frames, and environmental shape provides a salient cue to reference frame selection. To date, however, the environmental cues responsible for influencing reference frame selection remain relatively unknown. To connect research on reference frame selection with that on orientation via environmental shape, we explored the extent to which geometric cues were incidentally encoded and represented in memory by evaluating their influence on reference frame selection. Using a virtual environment equipped with a head-mounted-display, we presented participants with to-be-remembered object arrays. We manipulated whether the experienced viewpoint was aligned or misaligned with global (i.e., the principal axis of space) or local (i.e., wall orientations) geometric cues. During subsequent judgments of relative direction (i.e., participants imagined standing at one object, facing a second object, and pointed toward a third object), we show that performance was best when imagining perspectives aligned with these geometric cues; moreover, global geometric cues were sufficient for reference frame selection, global and local geometric cues were capable of exerting differential influence on reference frame selection, and performance from experienced-imagined perspectives was equivalent to novel-imagined perspectives aligned with geometric cues. These results explicitly connect theory regarding spatial reference frame selection and spatial orientation via environmental shape and indicate that spatial memories are organized around fundamental geometric properties of space.     

Prospective memory in children and chimpanzees

Prospective memory (PM) involves remembering to do something at a specific time in the future. Here, we investigate the beginnings of this ability in young children (3-year-olds; Homo sapiens) and chimpanzees (Pan troglodytes) using an analogous task. Subjects were given a choice between two toys (children) or two food items (chimpanzees). The selected item was delivered immediately, whereas the unselected item was hidden in an opaque container. After completing an ongoing quantity discrimination task, subjects could request the hidden item by asking for it (children) or by pointing to the container and identifying the item on a symbol board (chimpanzees). Children and chimpanzees showed evidence of prospective-like memory in this task, as evidenced by successful retrieval of the item at the end of the task, sometimes spontaneously with no prompting from the experimenter. These findings contribute to our understanding of PM from an ontogenetic and comparative perspective.     

Interference effects by spatial proximity and age-related declines in spatial memory by Japanese monkeys (Macaca fuscata): deficits in the combined use of multiple spatial cues

Spatial information processing was assessed in 3 young (4-10 years old) and 4 aged (24-25 years old) Japanese monkeys (Macaca fuscata) on 3 delayed nonmatching-to-position (DNMP) tests with relatively short delays of 5 s. Each test had 3 conditions of different horizontal distances between sample and to-be-nonmatched positions. Experiment 1 demonstrated that the performance on the DNMP test in both age groups was impaired when 2 stimulus positions were located next to each other; however, it was fairly accurate when they were located farther apart, suggesting that interference is introduced by spatial proximity. Experiment 2 revealed age-related differences in the situation in which an additional spatial cue, depth information, was available by extending the stimulus array of the DNMP test to a 4 x 2 matrix. In this test, young monkeys performed accurately irrespective of position distance between stimuli, whereas the aged monkeys' performance remained the same as before. Experiment 3 confirmed that the recognition ability in aged monkeys was well preserved on DNMP tests with different objects. These patterns of results indicate that the ability to use information from multiple spatial cues is not accessible to the aged monkeys.     

Spatio-temporal working-memory and short-term object-location tasks use different memory mechanisms

Spatial short-term memory for objects' locations was investigated in a spatial relocation task. During maintenance, dynamic visual noise or spatial tapping were administered as visual or spatial secondary tasks, respectively. Because memory for location should tap the visual component of working memory, a visual but not a spatial secondary task should impair location memory. In fact, neither of the tasks impaired memory (Experiment 1), although the expected dissociation between visual and spatial components was clearly confirmed for a spatio-temporal main task (Corsi test) (Experiment 2). We then contrasted location memory for pictures of objects and of nonsense figures under visual interference. Real objects were relocated much better than nonsense figures, and visual noise was again ineffective (Experiment 3). When spatial tapping was combined with the same material (Experiment 3a), again no influence on memory for locations of objects was observed and only a small influence on remembering nonsense figures. We suggest that the Corsi and the relocation VSWM-tasks use different memory mechanisms. The configuration of objects is reconstructed from perceptual records in an episodic buffer, provided by the same structures that enable visual memory after longer intervals. Rehearsal is not necessary for the persistence of these traces. In contrast, in the Corsi task remembering, a temporal sequence across homogeneous locations needs spatio-temporal marking and therefore active rehearsal of the locations by shifting spatial attention. A spatially demanding secondary task during retention interrupts this rehearsal.     

Environment size and the use of feature and geometric cues for reorientation

We tested associative-based accounts of orientation by investigating the influence of environment size on the use of feature and geometric cues for reorientation. Two groups of participants were trained in dynamic three-dimensional virtual rectangular environments that differed in size to find a distinctly colored bin located at one of the four corners. Subsequently, we probed the reliance on feature and geometric cues for reorientation during test trials by presenting six trial types: Small Geometry Only, Large Geometry Only, Small Cue Conflict, Large Cue Conflict, Small Distal, and Large Distal. During Geometry Only test trials, all bins were black; thus, all distinctive featural information was removed leaving only geometric cues. For Cue Conflict test trials, all colored bins were shifted counter-clockwise one corner; thus, the geometric cues from the trained corner and the trained color were in direct conflict. During Distal test trials, the bin in the geometrically incorrect corner farthest from the trained corner was colored the same as during training; the remaining three bins were black. Thus, only this distant feature cue could be used to determine the location of the goal bin. Results suggested that geometric cues were used across changes in environment size, featural cues exerted greater influence when in conflict with geometric cues, and the far featural cue was used to disambiguate the correct from the rotationally equivalent location. In short, both feature and geometric cues were used for reorientation, and environment size influenced the relative use of feature and geometric cues. Collectively, our results provide evidence against associative-based accounts of orientation.     

Virtual human analogs to rodent spatial pattern separation and completion memory tasks

Spatial pattern separation (SPS) and spatial pattern completion (SPC) have played an increasingly important role in computational and rodent literatures as processes underlying associative memory. SPS and SPC are complementary processes, allowing the formation of unique representations and the reconstruction of complete spatial environments based on partial spatial information. We present two novel computerized navigational tasks as human analogs of well-established rat SPS and SPC tasks. Results from these tasks show that human participants are sensitive to increasing SPS and SPC demands. Specifically, memory accuracy decreased with decreasing separation distance between target and foil locations in the SPS task and with decreasing number of distal spatial cues in the SPC task. These tasks set the stage for valuable future directions, including the use of these tasks with imaging and clinical populations.     

The use of recognition information and additional cues in inferences from memory

Goldstein and Gigerenzer's (2002) "Recognition Heuristic" (RH) was tested for its empirical validity in an experimental paradigm with induced recognition of objects. RH claims that upon inferring which of two objects (e.g., cities) scores higher on a criterion (e.g., city size), a recognized object will be chosen over an unrecognized one, if the recognition is a valid predictor of the criterion without considering additional object information. Trying to avoid potential shortcomings of former studies, we (a) used the city population task, (b) provided additional cue information only for recognized cities, and (c) had participants draw inferences from memory. Participants learned city names and additional information about some cities. They also learned that recognition and the additional information were valid predictors of the criterion "city size". In a subsequent decision phase, the additional information about the cities in memory strongly affected the inferences, suggesting that recognition information is clearly integrated into judgments, but by no means in a non-compensatory fashion that would dominate every other cue.     

The utilization of external and movement cues in simple spatial tasks by blind and sighted children.

The role of visual experience in coding spatial position by movements or by external cues was examined in simple (nonrotational) shift tasks with blind and sighted children. Age and the salience of external cues were also of interest. Results showed that the congenitally totally blind used movement cues significantly more even when external cues were present and prominent. The blind with minimal visual experience coded by external cues, but made errors beyond the age by which blindfolded sighted children performed correctly. It was argued that visual experience affects coding by drawing attention to external cues, and by providing more adequate spatial information than other sources usually available to the blind. In its absence, movement coding and self-reference can become preferred strategies.     

Variation in the use of cues to guide visual working memory

Across two experiments we examined individual differences in the use of precues and retrocues to guide the selective encoding and maintenance of information in visual working memory (WM). In Experiment 1, we used spatial cues to indicate which item would be tested either before or after the presentation of a memory array. Regression analyses allowed us to separate variance due to visual WM capacity and attention control differences. In Experiment 2, we used categorical cues to indicate from which subset of items the tested item would be drawn, and we measured attention control with two independent tasks. Collectively, the results supported the idea that an element of individual differences in WM is the ability to flexibly allocate attention to encode only relevant information and subsequently select relevant information during maintenance stages.     

Cross-domain interference costs during concurrent verbal and spatial serial memory tasks are asymmetric

Some evidence suggests that memory for serial order is domain-general. Evidence also points to asymmetries in interference between verbal and visual–spatial tasks. We confirm that concurrently remembering verbal and spatial serial lists provokes substantial interference compared with remembering a single list, but we further investigate the impact of this interference throughout the serial position curve, where asymmetries are indeed apparent. A concurrent verbal order memory task affects spatial memory performance throughout the serial positions of the list, but performing a spatial order task affects memory for the verbal serial list only for early list items; in the verbal task only, the final items are unaffected by a concurrent task. Adding suffixes eliminates this asymmetry, resulting in impairment throughout the list for both tasks. These results suggest that domain-general working memory resources may be supplemented with resources specific to the verbal domain, but perhaps not with equivalent spatial resources.