Changes in attention to relevant and irrelevant stimuli during spatial learning

Rats were trained in 2 experiments to find a submerged platform that was situated in 1 of 2 of the 4 corners of a rectangular pool with a curved long wall. Different landmarks occupied 2 of the corners on every trial, and the platform was always situated near a landmark. For the place group in each experiment, the location of the platform was indicated by the shape of the pool and stimuli outside the pool (place cues), but not the landmarks within the pool. For the landmark groups, the landmarks, not the place cues, indicated where the platform could be found. During Stage 2, 2 of the place cues were relevant, and 2 of the landmarks were irrelevant, for a new discrimination. The place cues better controlled searching for the platform in the place group than in the landmark group when the place cues had initially been relevant by signaling the presence (Experiment 1) or the absence (Experiment 2) of the platform. The results show that animals pay more attention to relevant than irrelevant cues.     

The effects of pre-exposure on escape from a Morris pool

In two experiments rats were pre-exposed to the landmarks surrounding a Morris pool while they swam to a platform with a beacon attached to it. They were then required to escape from the pool by finding the platform, without the beacon, in a new position. When the platform remained in the same place for each pre-exposure session, but was moved from session to session, then subsequent escape from the pool was more rapid than when the landmarks were not visible during pre-exposure (Experiment 1). But when the platform was moved from trial to trial during pre-exposure, then subsequent escape from the pool was disrupted (Experiments 1 and 2). It is proposed that pre-exposure to the landmark alters the attention that is paid to them, which then influences how readily the landmarks can be used to identify the new position of the platform.     

The influence of landmark salience in a navigation task: an additive effect between its components

In 2 experiments, rats were trained in a Morris pool to find a hidden platform in the presence of 1 landmark. After acquisition, the rats were tested without the platform. Experiment 1 tested whether the size of a landmark and its relative distance from the platform are additive effects. On test, the rats' best performance was with a near and big landmark; intermediate performance was with either a near and small landmark or a far and big one; and the worst performance was with a far and small landmark. The results of Experiment 2 suggested that the different distances from the goal of the 2 landmarks might not be sufficient to explain the previous results.     

Effect of absolute spatial proximity between a landmark and a goal

In two experiments rats were trained in a Morris pool to find a hidden platform in the presence of a single landmark. Circular black curtains surrounded the pool, with the single landmark inside this enclosure, so that no other room cues could provide additional information about the location of the platform. This landmark was hung from a false ceiling and rotated from trial to trial, and the position of the platform also changed on each trial, thus preserving a constant relation between the platform and the landmark. In Experiment 1, the position of the landmark was exactly above the hidden platform in Group Above and was relatively close to the hidden platform in Group Near. At the end of acquisition, test trials without the platform revealed a difference between the groups. Although a preference for searching in the correct quadrant of the pool, where the platform should have been, was found in both groups, this preference was significantly higher for Group Above. In Experiment 2, new rats in Group Near were compared to rats for which the position of the landmark was relatively far from the hidden platform in Group Far. Test trials revealed a preference for searching in the correct quadrant of the pool in both groups, but this preference was significantly higher for Group Near. The implication of these results is that the control acquired by a single landmark is different depending on its relative distance from a hidden platform: Closer landmarks acquire better control than distant ones. These results show a clear parallelism in comparison with the effect of absolute temporal proximity of the CS to the US in classical conditioning.     

Transfer of spatial behaviour controlled by a landmark array with a distinctive shape

In two experiments, rats swam to a submerged platform in one corner of a rectangular or kite-shaped array created by four identical landmarks attached to the walls of a circular pool. After training in the rectangular array, rats expressed a preference for the corner in the kite-shaped array that was geometrically equivalent to where the platform was located previously. After training in either array, the removal of two landmarks from the rectangular array, or the landmark at the apex of the kite-shaped array, did not affect the control over searching exerted by the remaining landmarks. The results imply that rats use local rather than global spatial representations when searching for a hidden goal with reference to an array of landmarks.     

Failure of a landmark to restrict spatial learning based on the shape of the environment

Rats were required to find a submerged platform in the corner of a swimming pool with a distinctive shape. A landmark near the platform did not interfere with the control acquired by the pool's shape over searching for the platform. This outcome was observed with an overshadowing and a blocking design. A comparison of the ease with which the landmark and the pool's shape gained control over searching for the platform indicates that the failure of overshadowing and blocking was not a consequence of the landmark being less salient than the shape of the pool. The results are not readily explained by theories of associative learning, but they are consistent with the claim that learning about the shape of the environment takes place in a dedicated module, which excludes information about the significance of individual landmarks.     

The influence of passive preexposure on escape from a Morris pool

In 3 experiments rats were preexposed to the landmarks that surround a Morris pool by being placed on a submerged platform within the pool. They were then required to escape from the pool by swimming to the platform, which was in a location that had not been used during preexposure. Preexposure facilitated subsequent escape from the pool, provided that the platform was not moved during preexposure and the relative position of the landmarks to each other remained constant throughout preexposure. In contrast, if during preexposure the platform was moved from session to session (Experiment 1), or the array of landmarks was altered unsystematically from trial to trial (Experiments 2 and 3), then subsequent learning to escape from the pool was disrupted. These findings suggest that the effects of preexposure to the landmarks in a Morris pool is determined by whether or not they are of relevance for identifying the location of the platform. When they are relevant, then subsequent learning is facilitated, but when they are irrelevant, then subsequent learning is disrupted.     

A computer simulation model of rats’ place navigation in the Morris water maze

Rats rapidly learn to swim from a variety of starting locations, including novel ones, to a small invisible platform submerged in a pool of cool opaque liquid. A computer simulation based on a simple perceptual memory-matching model successfully mimics this ability. The model assumes that the rat, when it successfully reaches the platform, notes the distance to prominent extramaze landmarks and stores this perceptual information in memory. When placed in the pool on subsequent trials, the simulated rat attempts to match perceived distance between itself and the landmarks to the remembered distances from the platform to the landmark. The model accounts for many of the known facts about rats’ behavior in the swimming pool and makes some interesting predictions that could be easily tested experimentally. The model has the advantage, relative to other cognitive map models, of specifying how spatial information is represented in memory and how this information guides behavior.     

Eye tracking, strategies, and sex differences in virtual navigation

Reports of sex differences in wayfinding have typically used paradigms sensitive to the female advantage (navigation by landmarks) or sensitive to the male advantage (navigation by cardinal directions, Euclidian coordinates, environmental geometry, and absolute distances). The current virtual navigation paradigm allowed both men and women an equal advantage. We studied sex differences by systematically varying the number of landmarks. Eye tracking was used to quantify sex differences in landmark utilisation as participants solved an eight-arm radial maze task within different virtual environments. To solve the task, participants were required to remember the locations of target objects within environments containing 0, 2, 4, 6, or 8 landmarks. We found that, as the number of landmarks available in the environment increases, the proportion of time men and women spend looking at landmarks and the number of landmarks they use to find their way increases. Eye tracking confirmed that women rely more on landmarks to navigate, although landmark fixations were also associated with an increase in task completion time. Sex differences in navigational behaviour occurred only in environments devoid of landmarks and disappeared in environments containing multiple landmarks. Moreover, women showed sustained landmark-oriented gaze, while men's decreased over time. Finally, we found that men and women use spatial and response strategies to the same extent. Together, these results shed new light on the discrepancy in landmark utilisation between men and women and help explain the differences in navigational behaviour previously reported.     

Disorientation inhibits landmark use in 12-18-month-old infants

Recent research has indicated that, particularly under conditions of inertial disorientation, mammals may be sensitive to landmark configuration geometry at the expense of individual featural information when locating hidden goals. The current study sought to establish whether landmark use could be demonstrated in 12-18-month-old infants with and without a disorientation procedure, and with geometrically ambiguous landmark configurations. A peekaboo paradigm was employed in which infants learned to anticipate a peekaboo event after a cue from two locations within a circular arena, followed by a test trial from a novel position in which no peekaboo occurred after the cue. In all conditions, an isosceles triangle arrangement of landmarks was used, with peekaboo occurring between the landmarks of one of the two equal "sides", thus being geometrically ambiguous. In two conditions, the landmarks were distinctive, and in two further conditions, they were identical. In one of the distinctive conditions and one of the identical landmark conditions, infants underwent a disorientation procedure in between training and test trials. Only oriented infants with distinctive landmarks were successful in test trials, thus suggesting that infants are able to use the individual features of landmarks to locate a goal, but can only do so if oriented.     

Generalization decrement and not overshadowing by associative competition among pairs of landmarks in a navigation task

When they are trained in a Morris water maze to find a hidden platform, whose location is defined by a number of equally spaced visual landmarks round the circumference of the pool, rats are equally able to find the platform when tested with any two of the landmarks (Prados, & Trobalon, 1998; Rodrigo, Chamizo, McLaren, & Mackintosh, 1997). This suggests that none of the landmarks was completely overshadowed by any of the others. In Experiment 1 one pair of groups was trained with four equally salient visual landmarks spaced at equal intervals around the edge of the pool, while a second pair was trained with two landmarks only, either relatively close to or far from the hidden platform. After extensive training, both male and female rats showed a reciprocal overshadowing effect: on a test with two landmarks only (either close to or far from the platform), rats trained with four landmarks spent less time in the platform quadrant than those trained with only two. Experiment 2 showed that animals trained with two landmarks and then tested with four also performed worse on test than those trained and tested with two landmarks only. This suggests that generalization decrement, rather than associative competition, provides a sufficient explanation for the overshadowing observed in Experiment 1. Experiment 3 provided a within-experiment replication of the results of Experiments 1 and 2. Finally, Experiment 4 showed that rats trained with a configuration of two landmarks learn their identity.     

Global and local spatial landmarks: their role during foraging by Columbian ground squirrels (Spermophilus columbianus)

Locating food and refuge is essential for an animal's survival. However, little is known how mammals navigate under natural conditions and cope with given environmental constraints. In a series of six experiments, I investigated landmark-based navigation in free-ranging Columbian ground squirrels (Spermophilus columbianus). Squirrels were trained individually to find a baited platform within an array of nine identical platforms and artificial landmarks set up on their territories. After animals learned the location of the food platform in the array, the position of the latter with respect to local artificial, local natural, and global landmarks was manipulated, and the animal's ability to find the food platform was tested. When only positions of local artificial landmarks were changed, squirrels located food with high accuracy. When the location of the array relative to global landmarks was altered, food-finding accuracy decreased but remained significant. In the absence of known global landmarks, the presence of a familiar route and natural local landmarks resulted in significant but not highly accurate performance. Squirrels likely relied on multiple types of cues when orienting towards a food platform. Local landmarks were used only as a secondary mechanism of navigation, and were not attended to when a familiar route and known global landmarks were present. This study provided insights into landmark use by a wild mammal in a natural situation, and it demonstrated that an array of platforms can be employed to investigate landmark-based navigation under such conditions.     

The development of landmarks in spatial memory: the role of differential experience

To assess how differential experience with objects in a spatial array might serve to establish relative landmarks within the array, first and fifth graders learned models of a town and farm under two conditions. In a homogeneous condition, all elements in the array were visited an equal number of times. In a landmark condition, the relative landmark status of an element was established by distributing the same total number of visits unequally across the elements. The effects of such landmarks on spatial memory were assessed both in reconstructions of the entire array and by pairwise distance estimations. The landmark condition led to a general improvement in spatial recall accuracy as well as providing a relative landmark within the array to help organize the space. The results suggest that different levels of experience (controlling for overall experience and object salience) can establish elements as relative landmarks in spatial memory. Although there was clear developmental improvement in spatial memory, the specific landmark effects were similar for both first and fifth graders.     

Clark's nutcrackers (Nucifraga columbiana) and the effects of goal--landmark distance on overshadowing

Three groups of Clark's nutcrackers (Nucifraga columbiana) were trained to find a goal location defined by an array of 4 landmarks that varied in goal--landmark distance. The arrays for each group differed in the distance of the closest landmark and contained goal--landmark distances that were common across groups, allowing for the examination of the effects of both relative and absolute goal--landmark distance on encoding of a landmark array. All 3 groups readily learned the task and were subsequently tested in probe tests with only single landmarks from the array available. Search error in tests with single landmarks was compared both within and across groups. Results demonstrated that both relative and absolute goal-landmark distances are important in spatial search.     

Solving small spaces: investigating the use of landmark cues in brown capuchins (Cebus apella)

Some researchers have recently argued that humans may be unusual among primates in preferring to use landmark information when reasoning about some kinds of spatial problems. Some have explained this phenomenon by positing that our species’ tendency to prefer landmarks stems from a human-unique trait: language. Here, we test this hypothesis—that preferring to use landmarks to solve such tasks is related to language ability—by exploring landmark use in a spatial task in one non-human primate, the brown capuchin monkey (Cebus apella). We presented our subjects with the rotational displacement task, in which subjects attempt to relocate a reward hidden within an array of hiding locations which are subsequently rotated to a new position. Over several experiments, we varied the availability and the salience of a landmark cue within the array. Specifically, we varied (1) visual access to the array during rotation, (2) the type of landmark, (3) the consistency of the landmark qualities, and (4) the amount of exposure to the landmark. Across Experiments 1 through 4, capuchins did not successfully use landmarks cues, suggesting that non-linguistic primates may not spontaneously use landmarks to solve some spatial problems, as in this case of a small-scale dynamic spatial task. Importantly, we also observed that capuchins demonstrated some capacity to learn to use landmarks in Experiment 4, suggesting that non-linguistic creatures may be able to use some landmarks cues in similar spatial tasks with extensive training.     

Absence of overshadowing and blocking between landmarks and the geometric cues provided by the shape of a test arena

In three experiments rats were required to escape from a pool of water by swimming to a submerged platform. The position of the platform was determined by the shape of the pool, which was either rectangular or triangular. A landmark that was located on the surface of the pool near the platform failed to overshadow (Experiment 1) or block (Experiment 2) learning about the position of the platform with reference to the shape of the pool. Experiment 3 revealed a similar outcome with cues outside the pool, which could be used, in addition to the shape of the pool, to identify the location of the platform. These findings imply that theories of learning that assume that stimuli must compete with each other for the control that they acquire may not apply to spatial learning based on the shape of the environment.     

Gender-Based Navigation Stereotype Improves Men’s Search for a Hidden Goal

While a general stereotype exists that men are better at navigating than women, experimental evidence indicates that men and women differ in their use of spatial strategies, and this preference determines gender-differences. When both environmental geometry and landmark cues are available, men appear to learn to navigate using both types of cues, while women show a preference for using landmarks. Using a computer-generated task, 80 undergraduate students from North-East England learned to navigate to a hidden goal. Activating the general navigation stereotype improved the performance of men, compared to the control condition, both when only geometric cues and only landmark cues were present (stereotype lift), suggesting that activating a general stereotype can affect tasks both with (geometry) and without (landmark) established gender-differences in preference. In addition, in the test trial (hidden goal removed) women who learned to navigate using only landmarks spent longer in the correct location of the hidden goal than those who learned to navigate using only geometry. In contrast, the opposite result was found for men, suggesting that when only one cue-type is available, gender-differences still occur, with women better able to navigate using landmarks than geometry, while men seemed to learn more about the location of the goal with reference to geometric than landmark cues.     

Strategies in landmark use by orangutans and human children

Landmark use has been demonstrated in a variety of organisms, yet the manner in which landmarks are encoded and subsequently used appears to vary between and sometimes within species, even when faced with identical landmark arrays. In the present experiments, orangutans and human children were shown a square array of identical landmarks and were trained to locate a hidden goal in the centre of the array. In Experiments 1 and 2, the search space appeared to be discrete, with white gridlines dividing up the space, and in Experiments 3a and 3b, the search space was uniformly coloured, making it appear continuous. In all experiments, following training, subjects were given a single expansion test, to determine their landmark strategy use, based on peak search activity. The orangutans appeared to use absolute directional vectors from individual landmarks, with peak search activities on the inner corners of the square array, and they used this strategy persistently. In contrast, human children showed two landmark-based strategies, absolute directional vectors and a relational or “middle” strategy, with the majority of children starting their search in the middle region. Although some children, especially young children, persistently used one strategy like the orangutans, many changed strategies when the original one failed to yield the hidden goal.     

Learned predictiveness training modulates biases towards using boundary or landmark cues during navigation

A number of navigational theories state that learning about landmark information should not interfere with learning about shape information provided by the boundary walls of an environment. A common test of such theories has been to assess whether landmark information will overshadow, or restrict, learning about shape information. Whilst a number of studies have shown that landmarks are not able to overshadow learning about shape information, some have shown that landmarks can, in fact, overshadow learning about shape information. Given the continued importance of theories that grant the shape information that is provided by the boundary of an environment a special status during learning, the experiments presented here were designed to assess whether the relative salience of shape and landmark information could account for the discrepant results of overshadowing studies. In Experiment 1, participants were first trained that either the landmarks within an arena (landmark-relevant), or the shape information provided by the boundary walls of an arena (shape-relevant), were relevant to finding a hidden goal. In a subsequent stage, when novel landmark and shape information were made relevant to finding the hidden goal, landmarks dominated behaviour for those given landmark-relevant training, whereas shape information dominated behaviour for those given shape-relevant training. Experiment 2, which was conducted without prior relevance training, revealed that the landmark cues, unconditionally, dominated behaviour in our task. The results of the present experiments, and the conflicting results from previous overshadowing experiments, are explained in terms of associative models that incorporate an attention variant.     

Single landmark learning in rats: Sex differences in a navigation task

In Experiments 1 and 2, rats were trained in a Morris pool to find a hidden platform located some distance away from a single landmark. Males learned to swim to the platform faster than females, but on test trials without the platform, males, unlike females, spent less time in the platform quadrant of the pool in the second half of each test trial than in the first. They also showed greater persistence in searching in the platform quadrant over a series of extinction trials. In Experiments 3a and 3b, the problem was made easier by locating the platform closer to the solitary landmark. Now males and females learned to swim to the platform equally rapidly, and both stopped searching in the platform quadrant in the second half of each test trial. Experiment 4 ruled out the possibility that males´ shorter latencies to find the platform in Experiment 2 were due to their swimming faster than females.