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.     

Blocking in the Morris swimming pool.

Four experiments demonstrate that spatial blocking is governed by the same principles that govern blocking in Pavlovian conditioning. In the 2nd stage of each experiment, rats escaped from a Morris swimming pool by swimming to a submerged platform with a beacon attached to it. Test trials were then conducted in the absence of the platform and the beacon to assess the extent to which subjects had learned about the position of the platform with reference to the room cues. For the 1st stage of their training, rats either swam to the platform and beacon in the presence of curtains that prevented the room cues from being seen (Experiments 1 & 2), or they swam to the platform and beacon that were moved from trial to trial (Experiments 3 & 4). In each experiment, learning about the room cues in the 2nd stage of the experiment was blocked by the presence of the beacon. This blocking effect was disrupted by changing the appearance of the beacon for the 2nd stage of training or by restricting the amount of exposure to the beacon during the 1st phase of training.     

Influence of a beacon on spatial learning based on the shape of the test environment.

In 5 experiments rats were required to escape from a triangular shaped pool by swimming to a submerged platform. The principal group of interest in each experiment received training with a beacon attached to the platform. The purpose of the experiments was to assess if the beacon overshadowed (Experiments 1-4) or blocked (Experiment 5) learning about the position of the platform with reference to the shape of the pool. The platform was located in the center of the pool for the first 2 experiments and in a corner for the remaining experiments. Although there was an overshadowing effect in Experiment 1, the remaining experiments failed to reveal any disruptive influence of the beacon on learning based on the shape of the pool. Moreover, in Experiments 3-5 there was an indication that the beacon facilitated such learning. The results suggest that spatial learning based on the shape of a test environment may not take place in the same way as that based on more discrete landmarks.     

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.     

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.     

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.     

Absence of an interaction between navigational strategies based on local and distal landmarks

In 3 experiments, rats were required to escape from a Morris pool by swimming to a submerged platform that was located at the apex of a notional, equilateral triangle with 2 different landmarks occupying the corners at the base. Training for 1 group was always conducted in view of the landmarks surrounding the pool and with the triangular array in a fixed orientation. Subjects could therefore identify the direction of the platform from a single landmark within the pool by reference to cues outside the pool or to the other landmark within the pool. Both strategies were used, and the results from additional groups revealed that the first of these strategies did not affect the acquisition of the second one.     

Development of place navigation in rats from weaning to puberty

Young hooded rats were trained to escape onto a hidden platform after swimming in a pool of opaque water. Subjects 21, 28, 35, 42, and 64 days of age on the first training day were given 28 trials on 5 consecutive days. Half of the rats were required to localize the platform in relation to external room cues only ("place only" condition) and the other half were helped by the presence of a visible cue on the platform ("cue + place" condition). A deficiency in place navigation was observed in the 21- and 28-day groups; they showed slow escape and took circuitous routes more often than older rats. This deficiency was related to a poor spatial bias toward the training position when the subjects were allowed to swim for 30 s in the absence of the platform, at the end of the 28-trial training period (probe trial). The 35-day group showed adult-like learning ability in both training conditions, but failed to show searching behavior during the probe trial after having been trained in the presence of the proximal cue. Only rats older than 40 days showed typical adult behavior such as swimming directly toward the platform from any starting position and localized searching around the absent platform's position during the probe trial, no matter what the training conditions were. These results suggest that central nervous system structures responsible for place learning in the rat are functional from around 32 days of age, but fail to trigger searching behavior following cued training before the sixth week.     

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.     

Vestibular stimulation disrupts acquisition of place navigation in the Morris water tank task

The significance of vestibular input for place navigation in the Morris water maze was examined in 11 hooded rats. A 5-min rotation (2 Hz) immediately preceding retrieval of an overtrained place navigation task prolonged escape latencies by about 10 s corresponding to circular swimming induced by postrotatory aftereffects, but did not otherwise interfere with target location. A 1-min rotation immediately following the acquisition trial in the working memory version of the water maze task did not deteriorate performance in the retrieval trial performed 5 min later. Two components of the acquisition trial, i.e., active search of the hidden target (up to 1 min) and latent learning during the 30-s stay on the platform, contribute almost equally to the formation of the working memory trace. A 1-min rotation immediately preceding the isolated platform learning component impaired subsequent retrieval but was ineffective when applied 3 min earlier. Latent learning was completely disrupted when the platform with the animal was rotated during the entire 30-s stay on the target platform or when the 30-s stay on the stationary platform was followed 0 or 3 min later by a 30-s rotation on the platform. The interfering effect of the latter procedure was suppressed by covering the platform with the animal by an opaque cylinder. It is concluded that vestibular cues are particularly important for the orientation of the animal in the gravitation field and for the estimation of the angles between vectors plotted from the animal toward external landmarks. Agreement between vestibular and visual signals is a prerequisite of efficient navigation.     

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.     

Generalization gradients in a navigation task with rats

In two experiments in a Morris pool, rats were trained to find a hidden platform which was located in a specific position in relation to two objects, B and F, which were presented together, one in front of the other. One object, B, was just above the platform (a beacon for the platform, the critical object) while the second object, F, was above the edge of the pool (the frame of reference). Then the rats received test trials, without the platform, in which B was presented in different positions in relation to F (i.e., in relation to its original position). In the two experiments the test results showed a generalization gradient as a function of the relative distance of the two objects: more time searching in the B segment, where the platform should have been, when B was in the original position (i.e., in front of F), which decreased symmetrically with distance of B from F. The present experiments show for the first time generalization gradients with rats across spatial locations when working with a navigation task.     

Role of context in perceptual learning in maze discriminations

Three experiments with rats in a maze examined the effects of pre-exposure to the relevant discriminative stimuli (rubber and sandpaper-covered maze arms) or the extra-maze context (the maze was surrounded either by black curtains or by variety of extra-maze landmarks) on the learning of a discrimination between rubber and sandpaper arms. In Experiment 1, pre-exposure to the extra-maze context facilitated subsequent discrimination learning. Experiments 2 and 3 showed that pre-exposure to rubber and sandpaper arms facilitated subsequent discrimination learning only when these cues were presented in the same context during pre-exposure and discriminative training. Taken together, the results are consistent with the hypothesis that a major cause of perceptual learning is the latent inhibition of stimuli or features common to the two discriminative stimuli, and that such latent inhibition may be disrupted by a radical change of context.     

An assessment of response, direction and place learning by rats in a water T-maze

Behavioral data suggest that distinguishable orientations may be necessary for place learning even when distal cues define different start points in the room and a unique goal location. We examined whether changes in orientation are also important in place learning and navigation in a water T-maze. In Experiment 1, rats were trained to locate a hidden platform and given a no-platform probe trial after 16 and 64 trials with the maze moved to a new position. Direction and response strategies were more prevalent than a place strategy. In Experiment 2, acquisition of place, response and direction strategies was assessed in a water T-maze that was moved between two locations during training. Rats were impaired on the place task when the maze was translated (moved to the L or R) but were successful when the maze was rotated across trials. These data are consistent with findings from appetitive tasks.     

Navigation in the Morris swim task as a baseline for drug discrimination: a demonstration with morphine

A morphine versus saline discrimination was demonstrated using the Morris swim task as the behavioral baseline. The apparatus was a large circular pool filled with water made opaque by floating polypropylene pellets. Rats were placed in the tank in randomly selected locations (12 trials per session) and could escape by swimming to a platform submerged 2 cm below the surface. Morphine (5.6 mg/kg) or saline was injected prior to training sessions. The position of the platform in a given session depended on the drug condition, thus forming the basis for discriminative responding. Three of the 4 rats acquired the discrimination, as evidenced by direct swims to the condition-appropriate platform. Generalization probe sessions were conducted following acquisition. Probe sessions were preceded by injections of morphine (0, 1.0, 3.0, 5.6, or 10.0 mg/kg) and involved placing the rat in the pool for 1 min without a platform. Swim patterns revealed a gradient, with probe swimming more concentrated in the area of the morphine platform position after higher morphine doses. In addition, dose-dependent increases in the likelihood of swimming first to the morphine-associated platform location were obtained. These results illustrate the generality of drug discrimination across different behavioral procedures, and of particular interest with respect to spatial learning, demonstrate interoceptive stimulus control of navigation.     

Latent spatial learning in an environment with a distinctive shape

Four experiments were conducted with rats in order to determine whether being placed on a platform in one corner of a rectangular swimming pool results in latent spatial learning. Rats in Experiments 1-3 received four trials a day of being placed on the platform. During a subsequent test trial, in which they were released into the pool without the platform, the rats exhibited a preference for swimming in the correct corners of the pool (those with the same geometric properties as the corner containing the platform during training), than the two remaining, incorrect corners. This effect was seen when the interval between the final placement trial and the test trial was as much as 24 hr (Experiment 2) and after varying numbers of sessions of placement training (Experiment 3). Experiment 4 revealed that when the test took place in a kite-shaped arena, after placement training in a rectangle, a stronger preference was shown for the corner that was geometrically equivalent to the correct rather than the incorrect corners in the rectangle. The placement treatment is said to result in latent spatial learning based on the development of S-S associations.     

Does nondirectional signalization of target distance contribute to navigation in the Morris water maze?

The possibility that rats can navigate in the Morris water maze by reducing the difference between the memorized platform scene and the current sensory input was tested in nine blind rats. A computerized videosystem monitored the rats' movements in the pool and converted the rat-target distance into tones the frequency of which increased in 64 equal steps from 120 Hz at 128 cm to 7680 Hz at 0 cm. During 15 days of training to find a fixed platform position from different starting points (12 trials per day) average escape latencies decreased from 39.0 to 25.4 s. The performance significantly deteriorated when the acoustic distance signalization was omitted and/or when the target position was changed form trial to trial. It is concluded that blind rats solved the task by simultaneously employing search strategy based on position responses, mapping using acoustic background beacons, and distance reduction navigation. It is argued that the various strategies are additive and that their relative significance depends of the conditions of the experiment.     

Directional learning,but no spatial mapping by rats performing a navigational task in an inverted orientation

Previous studies have identified neurons throughout the rat limbic system that fire as a function of the animal’s head direction (HD). This HD signal is particularly robust when rats locomote in the horizontal and vertical planes, but is severely attenuated when locomoting upside-down (Calton & Taube, 2005). Given the hypothesis that the HD signal represents an animal’s sense of directional heading, we evaluated whether rats could accurately navigate in an inverted (upside-down) orientation. The task required the animals to find an escape hole while locomoting inverted on a circular platform suspended from the ceiling. In Experiment 1, Long-Evans rats were trained to navigate to the escape hole by locomoting from either one or four start points. Interestingly, no animals from the 4-start point group reached criterion, even after 29 days of training. Animals in the 1-start point group reached criterion after about six training sessions. In Experiment 2, probe tests revealed that animals navigating from either 1- or 2-start points utilized distal visual landmarks for accurate orientation. However, subsequent probe tests revealed that their performance was markedly attenuated when navigating to the escape hole from a novel start point. This absence of flexibility while navigating upside-down was confirmed in Experiment 3 where we show that the rats do not learn to reach a place, but instead learn separate trajectories to the target hole(s). Based on these results we argue that inverted navigation primarily involves a simple directional strategy based on visual landmarks.     

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.     

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.