A consistent but non-coincident visual pattern facilitates the learning of spatial relations among locations

Human participants searched in a dynamic three-dimensional computer-generated virtual-environment open-field search task for four hidden goal locations arranged in a diamond configuration located in a 5 × 5 matrix of raised bins. Participants were randomly assigned to one of two groups: visual pattern or visual random. All participants experienced 30 trials in which four goal locations maintained the same spatial relations to each other (i.e., a diamond pattern), but this diamond pattern moved to random locations within the 5 × 5 matrix from trial to trial. For participants in the visual pattern group, four locations were marked in a distinct color and arranged in a diamond pattern that moved to a random location independent of the hidden spatial pattern from trial to trial throughout the experimental session. For participants in the visual random group, four random locations were marked with a distinct color and moved to random locations independent from the hidden spatial pattern from trial to trial throughout the experimental session. As a result, the visual cues for the visual pattern group were consistent but not coincident with the hidden spatial pattern, whereas the visual cues for the visual random group were neither consistent nor coincident with the hidden spatial pattern. Results indicated that participants in both groups learned the spatial configuration of goal locations and that the presence of consistent but noncoincident visual cues facilitated the learning of spatial relations among locations.     

Facilitation of learning spatial relations among locations by visual cues: Implications for theoretical accounts of spatial learning

Human participants searched in a real environment or interactive 3-D virtual environment open field for four hidden goal locations arranged in a 2 × 2 square configuration in a 5 × 5 matrix of raised bins. The participants were randomly assigned to one of two groups: cues 1 pattern or pattern only. The participants experienced a training phase, followed by a testing phase. Visual cues specified the goal locations during training only for the cues 1 pattern group. Both groups were then tested in the absence of visual cues. The results in both environments indicated that the participants learned the spatial relations among goal locations. However, visual cues during training facilitated learning of the spatial relations among goal locations: In both environments, the participants trained with the visual cues made fewer errors during testing than did those trained only with the pattern. The results suggest that learning based on the spatial relations among locations may not be susceptible to cue competition effects and have implications for standard associative and dual-system accounts of spatial learning.     

Visual landmarks facilitate rodent spatial navigation in virtual reality environments

Because many different sensory modalities contribute to spatial learning in rodents, it has been difficult to determine whether spatial navigation can be guided solely by visual cues. Rodents moving within physical environments with visual cues engage a variety of nonvisual sensory systems that cannot be easily inhibited without lesioning brain areas. Virtual reality offers a unique approach to ask whether visual landmark cues alone are sufficient to improve performance in a spatial task. We found that mice could learn to navigate between two water reward locations along a virtual bidirectional linear track using a spherical treadmill. Mice exposed to a virtual environment with vivid visual cues rendered on a single monitor increased their performance over a 3-d training regimen. Training significantly increased the percentage of time avatars controlled by the mice spent near reward locations in probe trials without water rewards. Neither improvement during training or spatial learning for reward locations occurred with mice operating a virtual environment without vivid landmarks or with mice deprived of all visual feedback. Mice operating the vivid environment developed stereotyped avatar turning behaviors when alternating between reward zones that were positively correlated with their performance on the probe trial. These results suggest that mice are able to learn to navigate to specific locations using only visual cues presented within a virtual environment rendered on a single computer monitor.     

No evidence for overshadowing or facilitation of spatial pattern learning by visual cues

Two experiments were conducted to examine the effects of redundant and relevant visual cues on spatial pattern learning. Rats searched for hidden food items on the tops of poles that formed a square (Experiment 1) or a checkerboard (Experiment 2) pattern. The experimental groups were trained with visual cues that specified the locations of the baited poles. All groups were tested without visual cues so that any overshadowing or facilitation of spatial pattern learning by visual cues could be detected. Spatial choices were controlled by the spatial pattern and by the visual cues in both experiments. However, there was no evidence of overshadowing or facilitation of spatial pattern learning by visual cues in either experiment. The results are consistent with the idea that the representation of the spatial pattern that guides choices is not controlled by the same learning processes as those that produce associations between visual cues and food locations.     

Encoding of variability of landmark-based spatial information

Recent evidence suggests humans optimally weight visual and haptic information (i.e., in inverse proportion to their variances). A more recent proposal is that spatial information (i.e., distance and direction) may also adhere to Bayesian principles and be weighted in an optimal fashion. A fundamental assumption of this proposal is that participants encode the variability of spatial information. In a three-dimensional virtual-environment open-field search task, we provide evidence that participants encoded the variability of landmark-based spatial information. Specifically, participants searched for a hidden goal location in a 5 × 5 matrix of raised bins. Participants experienced five training phases in which they searched for a hidden goal that maintained a unique spatial relationship to each of four distinct landmarks. Each landmark was assigned an a priori value of locational uncertainty such that each varied in its ability to predict a goal (i.e., varied in number of potential goal locations). Following training, participants experienced conflict trials in which two distinct landmarks were presented simultaneously. Participants preferentially responded to the landmark with the lower uncertainty value (i.e., smaller number of potential goal locations). Results provide empirical evidence for the encoding of variability of landmark-based spatial information and have implications for theoretical accounts of spatial learning.     

Learning directions of objects specified by vision,spatial audition,or auditory spatial language

The modality by which object azimuths (directions) are presented affects learning of multiple locations. In Experiment 1, participants learned sets of three and five object azimuths specified by a visual virtual environment, spatial audition (3D sound), or auditory spatial language. Five azimuths were learned faster when specified by spatial modalities (vision, audition) than by language. Experiment 2 equated the modalities for proprioceptive cues and eliminated spatial cues unique to vision (optic flow) and audition (differential binaural signals). There remained a learning disadvantage for spatial language. We attribute this result to the cost of indirect processing from words to spatial representations.     

Multiple spatial learning strategies in goldfish (Carassius auratus)

There is a considerable amount of evidence that mammals and birds can use different spatial learning strategies based on multiple learning and memory systems. Unfortunately, only a few studies have investigated spatial learning and memory mechanisms in other vertebrates. This study aimed to identify the strategies used by goldfish to solve two different spatial tasks in a series of three experiments. In experiment 1, two groups of goldfish (Carassius auratus) were trained either in a spatial constancy task (SC), in which visual cues signalled the goal indirectly, or in a directly cued task (DC) in which similar cues signalled the goal directly. Transfer tests were conducted to study the effects of discrete cue deletion on the performance in both tasks. In these transfer tests the performance of the animals trained in the DC task dropped to chance level when the cue that signalled the goal directly was removed. In contrast, the removal of any single cue did not disrupt SC performance. In experiment 2, fish trained in the SC or the DC task were trained with the goal reversed. Goldfish in the SC group needed fewer sessions to master the reversal task than DC animals. Finally, experiment 3 investigated the effects of a substantial modification of the geometrical features of the apparatus on the performance of animals trained in the SC or in the DC condition. The performance of DC goldfish was not affected, whereas the same change disrupted performance in the SC animals despite the presence of the visual cues. These results suggest that there are separate spatial learning and memory systems in fish. Whereas the DC animals used a typical guidance strategy, relying only on the cue that signalled the goal directly, SC fish relied on a strategy with the properties of an actual spatial mapping system. Thus, the comparative approach points to the generality of these learning strategies among vertebrates. Received: 10 October 1998 / Accepted after revision: 16 April 1999     

Effects of spatial configurations on visual change detection: An account of bias changes

In order to determine whether people encode spatial configuration information when encoding visual displays, in four experiments, we investigated whether changes in task-irrelevant spatial configuration information would influence color change detection accuracy. In a change detection task, when objects in the test display were presented in new random locations, rather than identical or different locations preserving the overall configuration, participants were more likely to report that the colors had changed. This consistent bias across four experiments suggested that people encode task-irrelevant spatial configuration along with object information. Experiment 4 also demonstrated that only a low-false-alarm group of participants effectively bound spatial configuration information to object information, suggesting that these types of binding processes are open to strategic influences.     

Cross-sensory transfer of reference frames in spatial memory

Two experiments investigated whether visual cues influence spatial reference frame selection for locations learned through touch. Participants experienced visual cues emphasizing specific environmental axes and later learned objects through touch. Visual cues were manipulated and haptic learning conditions were held constant. Imagined perspective taking when recalling touched objects was best from perspectives aligned with visually-defined axes, providing evidence for cross-sensory reference frame transfer. These findings advance spatial memory theory by demonstrating that multimodal spatial information can be integrated within a common spatial representation.     

A viewpoint-independent process for spatial reorientation

Reorientation tasks, in which disoriented participants attempt to relocate objects using different visual cues, have previously been understood to depend on representing aspects of the global organisation of the space, for example its major axis for judgements based on geometry. Careful analysis of the visual information available for these tasks shows that successful performance could be based on the much simpler process of storing a visual ‘snapshot’ at the target location, and subsequently moving in order to match it. We tested 4-8-year olds on a new spatial reorientation task that could not be solved based on information directly contained in any retinal projection that they had been exposed to, but required participants to infer how the space is structured. Only 6-8-year olds showed flexible recall from novel viewpoints. Five-year olds were able to recall locations given movement information or a unique proximal landmark, but without these they could not do so, even when they were not disoriented or when the landmark was a familiar object. These results indicate that early developing spatial abilities based on view matching and self motion are supplemented by a later-developing process that takes into account the structure of spatial layouts and so enables flexible recall from arbitrary viewpoints.     

Implicitly learned suppression of irrelevant spatial locations

How do we ignore a salient, irrelevant stimulus whose location is predictable? A variety of studies using instructional manipulations have shown that participants possess the capacity to exert location-based suppression. However, for the visual search challenges we face in daily life, we are not often provided explicit instructions and are unlikely to consciously deliberate on what our best strategy might be. Instead, we might rely on our past experience—in the form of implicit learning—to exert strategic control. In this paper, we tested whether implicit learning could drive spatial suppression. In Experiment 1, participants searched displays in which one location contained a target, while another contained a salient distractor. An arrow cue pointed to the target location with 70 % validity. Also, unbeknownst to the participants, the same arrow cue predicted the distractor location with 70 % validity. Results showed facilitated RTs to the predicted target location, confirming target enhancement. Critically, distractor interference was reduced at the predicted distractor location, revealing that participants used spatial suppression. Further, we found that participants had no explicit knowledge of the cue-distractor contingencies, confirming that the learning was implicit. In Experiment 2, to seek further evidence for suppression, we modified the task to include occasional masked probes following the arrow cue; we found worse probe identification accuracy at the predicted distractor location than control locations, providing converging evidence that observers spatially suppressed the predicted distractor locations. These results reveal an ecologically desirable mechanism of suppression, which functions without the need for conscious knowledge or externally guided instructions.     

Allocentric spatial memory in humans with hippocampal lesions

An immersive virtual reality (IVR) system was used to investigate allocentric spatial memory in a patient (PR) who had selective hippocampal damage, and also in patients who had undergone unilateral temporal lobectomies (17 right TL and 19 left TL), their performance compared against normal control groups. A human analogue of the Olton [Olton (1979). Hippocampus, space, and memory. Behavioural Brain Science, 2, 315] spatial maze was developed, consisting of a virtual room, a central virtual circular table and an array of radially arranged up-turned 'shells.' The participant had to search these shells in turn in order to find a blue 'cube' that would then 'move' to another location and so on, until all the shells had been target locations. Within-search errors could be made when the participants returned to a previously visited location during a search, and between-search errors when they revisited previously successful, but now incorrect locations. PR made significantly more between-search errors than his control group, but showed no increase in within-search errors. The right TL group showed a similar pattern of impairment, but the left TL group showed no impairment. This finding implicates the right hippocampal formation in spatial memory functioning in a scenario in which the visual environment was controlled so as to eliminate extraneous visual cues.     

Considering spatial ability in virtual route learning in early aging

The aim of this study is to broaden our understanding of the construction and early decline of spatial mental representations in route learning, considering the extent to which spatial ability and age-related differences in environment learning interact. The experiment examines spatial mental representation derived from taking a realistic route acquired using virtual environment and compares individuals different in age but with similar spatial ability. A sample of 34 young (20–30 years) and 30 middle-aged (50–60 years) females with good mental rotation ability were chosen. Participants learned a complex route through its presentation in a virtual environment and then performed a series of tasks (landmark recognition, location of landmarks and verification of spatial relations). Results show that the two participant age groups had similar performance in landmark recognition task and in verification of sentences describing direct spatial relations; instead, the middle-aged group showed a poorer performance than younger in their ability to locate landmarks and to judge the truth of indirect spatial sentences. These results first suggest that spatial abilities have to be seriously considered to avoid any confusion with age, as age-related differences are attenuated when individuals are different in age but similar in spatial ability. Second they confirm a specific difficulty of older participants to handle spatial information in a global configuration.     

Development of spatial memory and spatial orientation in preschoolers and primary school children

The present study addresses the question of what kind of information children use when orientating in new environments, if given proximal and distal landmarks, and how spatial memory develops in the investigated age groups. Ten 5-year-old, ten 7-year-old and ten 10-year-old children were presented with the ‘Kiel Locomotor Maze’, containing features of the Radial Arm Maze and the Morris Water Maze, in order to assess spatial memory and orientation. Children had to learn to approach baited locations only. Task difficulty was equated with respect to the children's age. Training was given until the children reached criterion. During testing, the maze configuration and response requirements were systematically altered, including response rotation, cue rotation, cue deletion and response rotation with cue deletion in order to assess the spatial strategies used by the children. During training and testing, working-memory errors (WM), reference-memory errors (RM) and working-reference memory errors (WR) were recorded. As expected, no difference between age groups appeared during training, thus confirming comparable task difficulty across age groups. During testing, age groups differed significantly with regard to the orientation strategy used. The 5-year-olds were bound to a cue strategy, orientating towards local, proximal cues. The 10-year-olds mastered all tasks, thus displaying a place strategy, being able to use distal cues for orientation, and were even able to do so after being rotated 180°. The 7-year-olds proved to be at an age of transition: five of them were bound to a cue strategy, five children were able to adopt a place strategy. The differences in the orientation strategies used by children of different age groups was reflected by the sum of errors they made, also by RM. WM were found to be rare, especially in older children. We conclude that preschoolers use a cue strategy, that the development of place strategies occurs during primary school age and seems to be complete by the age of 10 years.     

Place and direction learning in a spatial T-maze task by neonatal piglets

Pigs are a valuable animal model for studying neurodevelopment in humans due to similarities in brain structure and growth. The development and validation of behavioral tests to assess learning and memory in neonatal piglets are needed. The present study evaluated the capability of 2-week old piglets to acquire a novel place and direction learning spatial T-maze task. Validity of the task was assessed by the administration of scopolamine, an anti-cholinergic drug that acts on the hippocampus and other related structures, to impair spatial memory. During acquisition, piglets were trained to locate a milk reward in a constant place in space, as well as direction (east or west), in a plus-shaped maze using extra-maze visual cues. Following acquisition, reward location was reversed and piglets were re-tested to assess learning and working memory. The performance of control piglets in the maze improved over time (P < 0.0001), reaching performance criterion (80 % correct) on day 5 of acquisition. Correct choices decreased in the reversal phase (P < 0.0001), but improved over time. In a separate study, piglets were injected daily with either phosphate-buffered saline (PBS; control) or scopolamine prior to testing. Piglets administered scopolamine showed impaired performance in the maze compared to controls (P = 0.03), failing to reach performance criterion after 6 days of acquisition testing. Collectively, these data demonstrate that neonatal piglets can be tested in a spatial T-maze task to assess hippocampal-dependent learning and memory.     

Hierarchical structures: chunking by food type facilitates spatial memory

Three experiments assessed the ability of male Sprague-Dawley rats to organize the spatial locations of different food types in a hierarchical manner to maximize the efficiency of working memory. Independent groups were exposed, on a 12-arm radial maze, to baiting arrangements varying in the stability of the pattern and type of food used as bait. Training rats with stable, differentiable baiting arrangements produced increased accuracy in choice performance, hierarchically ordered patterns of choice selection, slower growth of proactive interference when trials were massed, and the learning of the geometrical relations among food types independent of other extramaze cues. Such findings are strong evidence of the rat's ability to encode and use local cues for navigation, based on properties of the reinforcer. The application of a chunking strategy may provide for more efficient use of working memory by facilitating information storage, recall, or resetting mechanisms.     

Path information effects in visual and proprioceptive spatial learning

Objects in an environment are often encountered sequentially during spatial learning, forming a path along which object locations are experienced. The present study investigated the effect of spatial information conveyed through the path in visual and proprioceptive learning of a room-sized spatial layout, exploring whether different modalities differentially depend on the integrity of the path. Learning object locations along a coherent path was compared with learning them in a spatially random manner. Path integrity had little effect on visual learning, whereas learning with the coherent path produced better memory performance than random order learning for proprioceptive learning. These results suggest that path information has differential effects in visual and proprioceptive spatial learning, perhaps due to a difference in the way one establishes a reference frame for representing relative locations of objects.     

Forgetting in spatial memories acquired in a virtual environment

This study examined forgetting in spatial memories acquired in a virtual environment. In the two experiments, participants learned the locations of eight objects. In Experiment 1, the objects were presented as photographs in either a laboratory or in an equivalent virtual environment. Irrespective of learning condition, accuracy of recall of the locations was found to deteriorate after a retention interval of approximately 1 week. In Experiment 2, following virtual learning, three groups of participants performed a series of non‐spatial tasks of low, intermediate or high difficulty. The tasks were presented in a retention interval of 2 hours. A comparison of recall accuracy before and after presentation of the interference tasks indicated that that the groups were not differentially affected by the difficulty of the retroactive interference tasks. However, the groups differed in their subjective assessment of the mental workload involved in the tasks. The results are discussed with reference to a prominent theory of forgetting. Copyright © 2007 John Wiley & Sons, Ltd.     

Textures shape the attentional focus: Evidence from exogenous and endogenous cueing

The spatial cueing paradigm (Posner Quarterly Journal of Experimental Psychology 32:3–25, 1980) has often been used to investigate the time course of the deployment of visual attention in space. In a series of eight experiments we investigated whether spatial cues would not only enhance processing of stimuli presented at cued locations, but also enhance processing of the entire texture in which the stimuli were presented. Results showed highest accuracy for responses to stimuli presented at cued locations, a replication of the traditional cueing effect (Posner 1980). Additionally, stimuli presented at uncued locations were responded to with higher accuracy when they were presented inside the same texture as the cued location, as compared with stimuli presented outside the texture with the cued location. To investigate this texture advantage for both automatic and voluntary attention deployment, exogenous and endogenous cues were used. The texture advantage was observed for short interstimulus intervals (ISIs) of 50 and 100 ms for exogenous cues and for a longer ISI of 200 ms for endogenous cues. These findings indicate that the arrangement of task-irrelevant visual stimuli also can have a large impact on the cueing effect. This suggests that visual spatial attention spreads texture-wise across the visual field. Control experiments revealed that the homogeneity within texture elements contributes most to the effect but that the texture advantage is a function of both orientation contrast at the texture border and homogeneity within texture elements.     

Head-centred meridian effect on auditory spatial attention orienting

Six experiments examined the issue of whether one single system or separate systems underlie visual and auditory orienting of spatial attention. When auditory targets were used, reaction times were slower on trials in which cued and target locations were at opposite sides of the vertical head-centred meridian than on trials in which cued and target locations were at opposite sides of the vertical visual meridian or were not separated by any meridian. The head-centred meridian effect for auditory stimuli was apparent when targets were cued by either visual (Experiments 2, 3, and 6) or auditory cues (Experiment 5). Also, the head-centred meridian effect was found when targets were delivered either through headphones (Experiments 2, 3, and 5) or external loudspeakers (Experiment 6). Conversely, participants showed a visual meridian effect when they were required to respond to visual targets (Experiment 4). These results strongly suggest that auditory and visual spatial attention systems are indeed separate, as far as endogenous orienting is concerned.