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.     

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.     

Preexposure effects in the spatial domain: Dissociation between latent inhibition and perceptual learning

In 4 experiments, rats were given intermixed or blocked preexposure to an array of landmarks that subsequently defined the location of a hidden goal in a Morris pool task. Previous research has shown that intermixed preexposure to pairs of adjacent landmarks retards learning whereas preexposure to individual landmarks facilitates subsequent learning (J. Prados, V. D. Chamizo, & N. J. Mackintosh, 1999). Accordingly, in Experiment 1, intermixed and blocked preexposure to pairs of adjacent landmarks was found to retard learning. In Experiment 2, however, a scheduling effect was found: Rats given intermixed preexposure to the individual landmarks learned faster than rats given blocked or no preexposure. Experiment 3 showed that intermixed (but not blocked) preexposure to pairs of landmarks resulted in a facilitatory effect when preexposure and test were carried out in different contexts. Experiment 4 replicated within a single experiment the main results observed in Experiments 1 and 3. This pattern of results suggests that intermixed preexposure engages learning processes other than latent inhibition that facilitate subsequent learning of the 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.     

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.     

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.     

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.     

Estrogen replacement in ovariectomized rats affects strategy selection in the Morris water maze

While estrogen enhances performance on some tasks of learning and memory, it has impairing or no effects on others. It has been proposed that estrogen differentially affects performance on various tasks of learning and memory by influencing the strategy used to solve a task. The goal of the present study was to determine if estrogen would influence strategy selection in the Morris water maze. Long-Evans rats were ovariectomized and implanted with Silastic capsules containing 25% estradiol diluted in cholesterol or 100% cholesterol. Rats were trained in a water maze task in which multiple strategies were available for use to locate a hidden escape platform that was moved to a new location for each set of four daily trials. During 10 days of acquisition trials, a visible floating landmark was always located in a static position relative to the hidden escape platform. Additionally, fixed extramaze cues visible to the animals surrounded the maze. Following acquisition, 2 days of probe trials were conducted in which the static landmark was removed. Estrogen replacement in ovariectomized rats resulted in impaired performance across 10 days of acquisition. Additionally, while removal of the visible landmark during the probe trials had no effect on the performance of the females receiving estrogen, it significantly disrupted performance of females receiving cholesterol treatment. These results indicate that estrogen replacement in ovariectomized rats biases an animal against using a landmark or static cue to aid in the location of a hidden escape platform in the water maze.     

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.     

Drug-preexposure effects in flavor-aversion learning: associative interference by conditioned environmental stimuli

Previous research has documented that exposure to a drug reduces the ability of the drug to support subsequent flavor-aversion learning. The four experiments reported here examined the hypothesis that this drug-preexposure effect is due to associative interference from environmental stimuli associated with the drug effects during preexposure. When distinctive environmental stimuli (confinement in a black compartment) were present during drug preexposure, these stimuli significantly disrupted subsequent flavor-aversion learning. Furthermore, flavor conditioning was not significantly disrupted when drug preexposure occurred in the absence of salient environmental stimuli or when the previously conditioned environmental stimuli were extinguished prior to flavor conditioning. It is significant, and in contrast to other published research, that flavor conditioning was not disrupted when the distinctive cues paired with the drug during preexposure were absent at the time of the flavor-drug pairing. These results are thus consistent with results from conventional studies of stimulus blocking and suggest that associative processes can play a major role in the drug-preexposure effect.     

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.     

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.     

Spatial and reversal learning in the Morris water maze are largely resistant to six hours of REM sleep deprivation following training

This first test of the role of REM (rapid eye movement) sleep in reversal spatial learning is also the first attempt to replicate a much cited pair of papers reporting that REM sleep deprivation impairs the consolidation of initial spatial learning in the Morris water maze. We hypothesized that REM sleep deprivation following training would impair both hippocampus-dependent spatial learning and learning a new target location within a familiar environment: reversal learning. A 6-d protocol was divided into the initial spatial learning phase (3.5 d) immediately followed by the reversal phase (2.5 d). During the 6 h following four or 12 training trials/day of initial or reversal learning phases, REM sleep was eliminated and non-REM sleep left intact using the multiple inverted flowerpot method. Contrary to our hypotheses, REM sleep deprivation during four or 12 trials/day of initial spatial or reversal learning did not affect training performance. However, some probe trial measures indicated REM sleep-deprivation-associated impairment in initial spatial learning with four trials/day and enhancement of subsequent reversal learning. In naive animals, REM sleep deprivation during normal initial spatial learning was followed by a lack of preference for the subsequent reversal platform location during the probe. Our findings contradict reports that REM sleep is essential for spatial learning in the Morris water maze and newly reveal that short periods of REM sleep deprivation do not impair concurrent reversal learning. Effects on subsequent reversal learning are consistent with the idea that REM sleep serves the consolidation of incompletely learned items.     

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.     

Latent learning in a swimming pool place task by rats: Evidence for the use of associative and not cognitive mapping processes

Using a latent learning paradigm, the experiments examine two hypotheses of how rats solve place navigation tasks. According to associative theory, experience with all relevant cues is required for accurate navigation. According to cognitive mapping theory, animals can generate novel trajectories from knowledge of spatial relations of objects in the environment. Rats that had been placed on a platform, which was submerged in a pool of opaque water and was moved each day, were tested later for their ability to find the platform using only surrounding room cues. One 30-sec exposure to a platform location was effective in improving performance. Improvement was greatest when tests were given within minutes of placement, but facilitation was obtained for as long as 4 hr. Improved performance was obtained as soon as rats acquired the procedural aspects of the task but did not increase with subsequent practice. Improved performance was also obtained when pre-trained rats were tested in a novel environment. Despite the advantage conferred by exposure to the target platform, test swims were not accurate, placement-induced improvement was not as great as that following a single swimming trial, and placement combined with a swim resulted in best performance. The results suggest that rats use associative learning processes rather than cognitive mapping to solve place problems in a swimming pool.     

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.     

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.     

Rats learn the relationship between responding and environmental events: An expansion of the learned helplessness hypothesis

Previous research has shown that preexposure to inescapable shock interferes with subsequent acquisition of escape responding, while pretraining with escapable shock facilitates subsequent acquisition of a different escape response. It has also been demonstrated that interference and facilitation persist when the aversive event is changed between the two phases of training. The present experiment extended these findings, showing generalized learning from an appetitive to an aversive situation. Six groups of rats received the following treatment in the presence of discriminative stimuli: One group was trained to nose press for food, a second to chain pull for food, and a third to chain pull to escape or avoid shock. Two groups received either signalled free food or inescapable shock, and a naive control group received no pretreatment. All groups were then tested in a nose-press escape-avoidance situation. The three groups with prior response training acquired responding most rapidly, and at the same rate. The naive controls acquired responding slowly, and the two groups with response-independent histories did not acquire responding during the 5 days of training. It was concluded that rats learn the relationship between responding and environmental events and that such learning strongly influences subsequent learning.     

Evidence of blocking with geometric cues in a virtual watermaze

Three computer based experiments, testing human participants in a non-immersive virtual watermaze task, used a blocking design to assess whether two sets of geometric cues would compete in a manner described by associative models of learning. In stage 1, participants were required to discriminate between visually distinct platforms. In stage 2, additional spatial information was provided by the shape or the color of the walls of the pool. In a test trial, the platforms were removed and the spatial knowledge acquired regarding the position of the platform was assessed. Experimental groups were compared against control groups which did not receive stage 1 training. The unique color of the correct platform, in Experiments 1 and 3, disrupted learning about the colored walls but not the geometry of the pool. In Experiment 2, the correct platform was identifiable from its position within the three platform array. Learning the relative position of the correct platform within the array disrupted learning about its position relative to the geometry of the pool, but not to the colored walls. The results suggest that learning the position of a goal in relation to the geometry of the environment can be blocked but only by an alternative geometric cue.