Landmark Stability: Further Studies Pointing to a Role in Spatial Learning

Two experiments were conducted to investigate the possible role of landmark stability in spatial learning. Rats were trained to search in a large arena for food hidden at a consistent distance and direction from either a single radially symmetric landmark or an array of two landmarks. We varied the relative degree to which the landmark array and/or the cues of the training context predicted the location of food, without varying the conditional probability of food being available given either cue. Experiment 1 used vestibular disorientation to ensure control of search location by experimenter-controlled cues. The results showed that making either a single landmark or a cluster of two adjacent landmarks the sole spatial predictor of reward location reduced the accuracy of search compared to a condition where both the landmark array and context cues were reliable spatial predictors. Varying global landmark stability had no effect when training was conducted using an array of two landmarks located some distance from each other. Context cues, when tested alone, triggered very little searching in appropriate locations, and the absolute magnitude of control over search was insufficient to account for the superiority of stable landmarks. The better learning with a stable landmark, and the dependence of this effect on the geometrical arrangement of landmarks, points to the conditions of spatial learning involving additional principles to those of simple associative conditioning. Experiment 2 examined landmark stability using a single landmark and fixed directional cues in the absence of vestibular disorientation. This also revealed a relative advantage of landmark stability, but animals with a landmark that moved from trial to trial did show some evidence of learning. Context cues when tested alone had minimal influence. Parametric manipulation of landmark stability offers a novel way of influencing spatial learning and thus understanding better the process through which egocentric representations of perceived space are transformed into allocentric representations of the real world. The purpose of this paper is to describe two experiments concerned with identifying the psychological processes of allocentric spatial learning. The results point to the idea that landmark stability is an important factor in spatial learning. Specifically, they reveal that whether or not a landmark will be used for the purpose of representing the location of another object (including hidden objects) is influenced by whether it is perceived as geometrically stable with respect to at least one other landmark and/or certain geometric features of the environment. This phenomenon is relevant to the application of associative learning principles to the spatial domain.     

Caudate nucleus and memory for egocentric localization

A large body of evidence suggests that the caudate nucleus (CN) plays a critical role in the processing of spatial localization information. Furthermore, evidence has begun to accumulate that the CN is involved in the processing of a very specific class of spatial cues, namely, egocentric cues (localization with reference to the organism). This is in contrast to allocentric localization, where an organism localizes on the basis of cues external to the organism. One would then predict that lesions to the CN should disrupt performance on any tasks that depend chiefly on egocentric spatial cues, while leaving performance on allocentric tasks intact. To test this prediction, two groups of rats were trained on two different egocentric memory tasks and two different allocentric memory tasks. Specifically, one group of rats was trained on an adjacent-arm (egocentric) and an 8-arm radial maze task (allocentric). A second group of rats was trained on a right-left discrimination (egocentric) and a place-learning task (allocentric). After training, both groups received bilateral lesions of the CN. Results showed that CN-lesioned animals were profoundly impaired on retention of the egocentric tasks. In sharp contrast to this, the same animals were not or were only transiently impaired or transiently affected on allocentric tasks. Sham-operated controls were either unimpaired or transiently affected on all tasks. These findings further support the idea that the CN plays a critical modulatory role in the processing of egocentric spatial and not allocentric spatial cues.     

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.     

Beacon training in a water maze can facilitate and compete with subsequent room cue learning in rats

In Stage 1 of 4 experiments in which rats completed a water-maze blocking procedure, experimental groups were trained to use a predictive beacon (hanging above, connected to, or displaced from the platform) to find a submerged escape platform in the presence of predictive or irrelevant background cues and in the presence or absence of irrelevant landmarks. In Stage 2, a fixed beacon, landmarks, and background cues all predicted the platform location. A Room Test (landmarks and background cues only) showed that Stage 1 training with a fixed hanging beacon or the moving displaced beacon facilitated Stage 2 learning of predictive room cues for experimental relative to control subjects. In contrast, Stage 1 training with a moving pole beacon interfered with Stage 2 learning about predictive room cues relative to controls, whereas training with a fixed pole or moving hanging beacon had no effect. We conclude that multiple spatial learning processes influence locating an escape platform in the water maze.     

Spatial strategy elaboration in egocentric and allocentric tasks following medial prefrontal cortex lesions in the rat

We evaluated the role of the medial prefrontal cortex (mPFC) in the elaboration of egocentric navigation strategies in a water maze (WM). Lesions of mPFC cell bodies was achieved in 21 rats using bilateral injections of ibotenic acid (IA); 13 control rats were injected with saline. After 17 days, rats had to learn an allocentric (using external cues: 10 lesioned, 7 saline rats) or an egocentric WM (using internal/kinetic cues: 10 lesioned, 6 saline rats) over six trials in a same session. The initial trajectory on the sixth trial was used as an index of the elaboration of a navigation strategy. In the egocentric test, lesioned rats were more rarely located in the target quadrant than control rats. No differences were found between lesioned and control rats in the allocentric test. These results show that lesions of the mPFC impairs the capacity to elaborate an egocentric navigation strategy.     

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.     

Landmark Discrimination Learning in the Dog

Allocentric spatial memory was studied in dogs of varying ages and sources using a landmark discrimination task. The primary goal of this study was to develop a protocol to test landmark discrimination learning in the dog. Using a modified version of a landmark test developed for use in monkeys, we successfully trained dogs to make a spatial discrimination on the basis of the position of a visual landmark relative to two identical discriminanda. Task performance decreased, however, as the distance between the landmark and the “discriminandum” was increased. A subgroup of these dogs was also tested on a delayed nonmatching to position spatial memory task (DNMP), which relies on egocentric spatial cues. These findings suggest that dogs can acquire both allocentric and egocentric spatial tasks. These data provide a useful tool for evaluating the ability of canines to use allocentric cues in spatial learning.     

The relationship between allocentric and egocentric frames of reference and categorical and coordinate spatial information processing

We report two experiments on the relationship between allocentric/egocentric frames of reference and categorical/coordinate spatial relations. Jager and Postma (2003) suggest two theoretical possibilities about their relationship: categorical judgements are better when combined with an allocentric reference frame and coordinate judgements with an egocentric reference frame (interaction hypothesis); allocentric/egocentric and categorical/coordinate form independent dimensions (independence hypothesis). Participants saw stimuli comprising two vertical bars (targets), one above and the other below a horizontal bar. They had to judge whether the targets appeared on the same side (categorical) or at the same distance (coordinate) with respect either to their body-midline (egocentric) or to the centre of the horizontal bar (allocentric). The results from Experiment 1 showed a facilitation in the allocentric and categorical conditions. In line with the independence hypothesis, no interaction effect emerged. To see whether the results were affected by the visual salience of the stimuli, in Experiment 2 the luminance of the horizontal bar was reduced. As a consequence, a significant interaction effect emerged indicating that categorical judgements were more accurate than coordinate ones, and especially so in the allocentric condition. Furthermore, egocentric judgements were as accurate as allocentric ones with a specific improvement when combined with coordinate spatial relations. The data from Experiment 2 showed that the visual salience of stimuli affected the relationship between allocentric/egocentric and categorical/coordinate dimensions. This suggests that the emergence of a selective interaction between the two dimensions may be modulated by the characteristics of the task.     

Pigeons encode absolute distance but relational direction from landmarks and walls

In recent studies, researchers have examined animals' use of absolute or relational distances in finding a hidden goal. When trained with an array of landmarks, most animals use the default strategy of searching at an absolute distance from 1 or more landmarks. In contrast, when trained in enclosures, animals often use the relationship among walls. In the present study, pigeons were trained to find the center of an array of landmarks or a set of short walls that did not block external cues. Expansion tests showed that both groups of pigeons primarily used an absolute distance strategy. However, on rotational tests, pigeons continued to search in the center of the array, suggesting that direction was learned in relation to array.     

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.     

The relationship between allocentric and egocentric frames of reference and categorical and coordinate spatial information processing

We report two experiments on the relationship between allocentric/egocentric frames of reference and categorical/coordinate spatial relations. Jager and Postma (2003) suggest two theoretical possibilities about their relationship: categorical judgements are better when combined with an allocentric reference frame and coordinate judgements with an egocentric reference frame (interaction hypothesis); allocentric/egocentric and categorical/coordinate form independent dimensions (independence hypothesis). Participants saw stimuli comprising two vertical bars (targets), one above and the other below a horizontal bar. They had to judge whether the targets appeared on the same side (categorical) or at the same distance (coordinate) with respect either to their body-midline (egocentric) or to the centre of the horizontal bar (allocentric). The results from Experiment 1 showed a facilitation in the allocentric and categorical conditions. In line with the independence hypothesis, no interaction effect emerged. To see whether the results were affected by the visual salience of the stimuli, in Experiment 2 the luminance of the horizontal bar was reduced. As a consequence, a significant interaction effect emerged indicating that categorical judgements were more accurate than coordinate ones, and especially so in the allocentric condition. Furthermore, egocentric judgements were as accurate as allocentric ones with a specific improvement when combined with coordinate spatial relations. The data from Experiment 2 showed that the visual salience of stimuli affected the relationship between allocentric/egocentric and categorical/coordinate dimensions. This suggests that the emergence of a selective interaction between the two dimensions may be modulated by the characteristics of the task.     

Spatial encoding of hidden objects in dogs (Canis familiaris)

The authors investigated the type of spatial information that controls domestic dogs' (Canis familiaris) search behavior in a situation in which they have to locate a spatial position where they saw an object move and disappear. In Experiments 1 and 2, the authors manipulated all local and global sources of allocentric spatial information surrounding the hiding location. The results revealed that dogs relied on an egocentric frame of reference. Experiment 3 showed that dogs also encoded allocentric information when egocentric information was irrelevant. The authors conclude that dogs simultaneously encode both egocentric and allocentric spatial information to locate a spatial position, but they primarily base their search behavior on an egocentric frame of reference. The authors discuss under which natural conditions dogs might use these 2 sources of spatial information and detail the nature of spatial egocentric information and the circumstances underlying its use by dogs.     

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.     

Developmental time course of the acquisition of sequential egocentric and allocentric navigation strategies

Navigation in a complex environment can rely on the use of different spatial strategies. We have focused on the employment of “allocentric” (i.e., encoding interrelationships among environmental cues, movements, and the location of the goal) and “sequential egocentric” (i.e., sequences of body turns associated with specific choice points) strategies during navigation. To investigate the developmental pattern of these two strategies in school-aged children, we used a virtual reality paradigm in which the spontaneous or imposed use of both strategies could be assessed. Our results showed an increase in spontaneous use of the allocentric strategy and also an increase in reliance on environmental landmarks with age. Although a majority of the children spontaneously used the sequential egocentric strategy, all age groups performed above chance when the allocentric strategy was imposed. Altogether, our findings suggest that young children are able to employ an allocentric strategy but that the nature of this allocentric strategy changes progressively in a complex cognitive representation between 5 and 10 years of age.     

Landmark-based search memory in the domestic dog (Canis familiaris)

Recent studies have suggested that any animal that relies on landmark-based search memory encodes and uses metric properties of space to navigate. So far, however, metric information provided by landmarks has been predominantly investigated in avian species. In the present study, I investigated whether the domestic dog (Canis familiaris), a mammalian species, encodes the distance and direction from landmarks. Dogs were trained to find a ball hidden next to an array of two distinct landmarks that remained at a constant location in a room from trial to trial. After training, the dogs were occasionally tested on unrewarded conditions where the array was either left in its usual position or shifted laterally, perpendicularly, or diagonally relative to the rear wall of the room. Although the dogs significantly shifted their search as a function of the displacement of the landmarks, they did not search at the predicted coordinates of the goal relative to the shift of the landmarks, suggesting that the global cues available in the testing room were also encoded and used by dogs to locate the position of the goal.     

Effects of varying the amount of preexposure to spatial cues on a subsequent navigation task

In each of two experiments rats were preexposed to four compound landmarks (AX, BX, CX, and DX) one at a time; they were then trained to find a submerged platform located in a fixed position in a swimming pool using these same landmarks. When the preexposure was SHORT (4 sessions) it facilitated subsequent learning (a perceptual learning effect), whereas when rats were given a LONG preexposure phase (8 sessions) this facilitatory effect disappeared. EXTRA-LONG preexposure (16 sessions) reversed the facilitatory effect-that is to say, we observed retarded learning. The results show that rats' ability to navigate towards an invisible goal is affected by the length of their preexposure to the spatial cues that signal the location of the goal. These data are consistent with an associative analysis of the swimming pool navigation task.     

Dissociations between vision for perception and vision for action depend on the relative availability of egocentric and allocentric information

In three experiments, we scrutinized the dissociation between perception and action, as reflected by the contributions of egocentric and allocentric information. In Experiment 1, participants stood at the base of a large-scale one-tailed version of a Müller-Lyer illusion (with a hoop) and either threw a beanbag to the endpoint of the shaft or verbally estimated the egocentric distance to that location. The results confirmed an effect of the illusion on verbal estimates, but not on throwing, providing evidence for a dissociation between perception and action. In Experiment 2, participants observed a two-tailed version of the Müller-Lyer illusion from a distance of 1.5 m and performed the same tasks as in Experiment 1, yet neither the typical illusion effects nor a dissociation became apparent. Experiment 3 was a replication of Experiment 1, with the difference that participants stood at a distance of 1.5 m from the base of the one-tailed illusion. The results indicated an illusion effect on both the verbal estimate task and the throwing task; hence, there was no dissociation between perception and action. The presence (Exp. 1) and absence (Exp. 3) of a dissociation between perception and action may indicate that dissociations are a function of the relative availability of egocentric and allocentric information. When distance estimates are purely egocentric, dissociations between perception and action occur. However, when egocentric distance estimates have a (complementary) exocentric component, the use of allocentric information is promoted, and dissociations between perception and action are reduced or absent.     

Dissociation of Hippocampal and Striatal Contributions to Spatial Navigation in the Water Maze

Two experiments were conducted to compare the effects of fornix/fimbria and caudate-putamen lesions in Long–Evans hooded rats (Rattus norvegicus) trained on two water maze tasks that differed in the type of spatial localization required for optimum solution. In Experiment 1, the lesioned rats and surgical controls were trained on the standard place task in the water maze (Morris, 1981) and given two postacquisition tests (a platform removal probe and platform relocation test). In Experiment 2, rats with similar lesions and control rats were trained on a modified cue navigation task. Fornix/fimbria lesions impaired a late stage of place task acquisition but did not impair acquisition of the cue task. Caudate-putamen lesions resulted in a severe place acquisition impairment and a transient cue acquisition impairment, both of which were characterized by an initial tendency to swim near the wall of the pool. Post-hoc analyses of the direction and angles of departure from the start points suggested that rats with fornix/fimbria lesions used non-allocentric spatial strategies to solve the place task. These rats also demonstrated a significantly weakened spatial bias for the former training quadrant on the platform removal probe and reduced flexibility in navigating to a novel platform location on the platform relocation test. In contrast, rats with caudate-putamen lesions showed a significant spatial bias for the former training quadrant but failed to cross the exact location within the quadrant where the platform was formerly positioned. The results suggest that the hippocampus mediates the allocentric spatial component of the water maze place task while the dorsomedial striatum may play an important role in the acquisition of the procedural aspects of both place and cue versions of the task.     

Where am I and how will I get there from here? A role for posterior parietal cortex in the integration of spatial information and route planning

The ability of an organism to accurately navigate from one place to another requires integration of multiple spatial constructs, including the determination of one's position and direction in space relative to allocentric landmarks, movement velocity, and the perceived location of the goal of the movement. In this review, we propose that while limbic areas are important for the sense of spatial orientation, the posterior parietal cortex is responsible for relating this sense with the location of a navigational goal and in formulating a plan to attain it. Hence, the posterior parietal cortex is important for the computation of the correct trajectory or route to be followed while navigating. Prefrontal and motor areas are subsequently responsible for executing the planned movement. Using this theory, we are able to bridge the gap between the rodent and primate literatures by suggesting that the allocentric role of the rodent PPC is largely analogous to the egocentric role typically emphasized in primates, that is, the integration of spatial orientation with potential goals in the planning of goal-directed movements.     

听障和听力正常人群空间主导性和空间参照框架的交互作用

通过要求被试分别在近处空间和远处空间完成空间参照框架的判断任务, 考察了听障和听力正常人群空间主导性和空间参照框架的交互作用。结果表明：(1)相对于听力正常人群, 听障人群完成自我参照框架判断任务的反应时更长, 而在完成环境参照框架判断任务无显著差异; (2)听障人群和听力正常人群空间主导性和空间参照框架交互作用呈现出相反模式。研究表明, 听障人群在听力功能受损后, 其空间主导性和空间参照框架的交互作用也产生了变化。