Perceptual learning in maze discriminations

In Experiment 1, rats were trained on a discrimination between rubber- and sandpaper-covered arms of a maze after one group had been pre-exposed to these intra-maze cues. Pre-exposure facilitated subsequent discrimination learning, unless the discrimination was made easier by adding further discriminative stimuli, when it now significantly retarded learning. In Experiment 2, rats were trained on an extra-maze spatial discrimination, again after one group, but not another, had been pre-exposed to the extra-maze landmarks. Here too, pre-exposure facilitated subsequent discrimination learning, unless the discrimination was made substantially easier by arranging that the two arms between which rats had to choose were always separated by 135°. The results of both experiments can be explained by supposing that perceptual learning depends on the presence of features common to S+ and S-.     

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.     

Blocking and overshadowing between intra-maze and extra-maze cues: A test of the independence of locale and guidance learning

Rats were trained on an elevated maze where the rewarded alternative was defined either in terms of intra-maze cues (rubber or sandpaper flooring on rewarded and unrewarded arms, regardless of their position) or in terms of extra-maze cues (the correct arm always pointed toward a particular corner of the room, and was sometimes covered with rubber and sometimes with sandpaper), or where both sets of cues were simultaneously relevant. In Experiment 1 rats pretrained with either intra-maze or extra-maze cues alone relevant learned less about the other set of cues than non-pretrained control groups, when, in a second phase of the experiment, both sets of cues were simultaneously relevant. Experiment 2 confirmed that intra-maze cues could block extra-maze cues, and ruled out one alternative explanation of the results of Experiment 1. Experiment 3 showed that extra-maze cues overshadowed intra-maze cues, but that there was no reciprocal overshadowing of extra-maze by intra-maze cues. This was despite the fact that animals learned the intra-maze discrimination significantly faster than the extra-maze discrimination. Experiment 3 also suggested that rats did not solve the extra-maze discrimination by learning to approach or avoid specific extra-maze cues, but rather by locating the correct arm by reference to the entire set of extra-maze cues. The results suggest that locale or place learning and cue or guidance learning, in O'Keefe and Nadel's (1978) terminology, interact with one another in much the same way as does learning about any pair of stimuli in a Pavlovian conditioning experiment.     

Reciprocal overshadowing between intra-maze and extra-maze cues

In three experiments, rats learned a maze discrimination where the location of food was defined either by reference to extra-maze cues alone, or by both extra- and intra-maze cues. Experiment 1 confirmed earlier results in showing that the presence of intra-maze cues failed to overshadow learning about extra-maze cues, in spite of the former's apparently greater salience. Experiment 2, however, suggested that this result was an artefactual consequence of differences between groups in the proportion of reinforced and unreinforced trials during the course of discriminative training. In Experiment 3, the discrimination was taught by a series of reinforced and unreinforced placement trials, and a significant overshadowing effect was observed. Intra-maze and extra-maze cues seem to compete for association with reinforcement in exactly the same way as any other cues.     

Pre-training to find a hidden platform in the Morris water maze can compensate for a deficit to find a cued platform in a rat model of Parkinson's disease

The bilateral intranigral infusion of 1 micromol 1-methyl-4-phenyl-1,2,3,6-tetrahydropyridine (MPTP) in adult male Wistar rats caused a specific and partial loss of substantia nigra pars compacta (SNc) dopamine neurons, a partial depletion of striatal dopamine, and a deficit to learn the intra-maze cued version of the Morris water maze. Pre-training the SNc rats in the spatial version of the water maze or simply maintaining the animals on the water maze platform reversed this deficit. This improvement was even observed when the order of the extra-maze cues presented to the rats during pre-training of the spatial version was changed during training of the intra-maze cued version. However, this deficit was not reversed either by maintaining the animals on the platform if the spatial cues were surrounded and covered with a curtain or by swimming sessions in the maze without the escape platform and the curtain. These findings suggest that none of the following elements alone, learned during the spatial task pre-training, could help SNc rats learn the intra-maze cued task: improvement of swimming skills or knowledge of the existence of the escape platform; distance between the platform and the border of the pool; location of a particular extra-maze cue; relations among extra-maze cues. However, the simultaneous presence of the escape platform and extra-maze cues (irrespective of their relational configuration) during the pre-training sessions proved to be necessary for this improving effect to occur.     

Effects of pre-exposure to the same or different pattern of extra-maze cues on subsequent extra-maze discrimination

In two experiments, rats learned a spatial discrimination between maze arms defined by their relationship to a variety of extra-maze cues. Prior exposure to the actual arms between which animals were required to discriminate tended to retard subsequent learning (by comparison with a control group either given no pre-exposure to the extra-maze cues or exposed only to arms pointing in the opposite direction), whereas prior exposure to arms intermediate between those used in discrimination training tended to facilitate subsequent learning. These results are consistent with the suggestion that pre-exposure will facilitate discrimination learning when it reduces the associability of features or elements common to the stimuli between which animals are required to discriminate, more than it reduces the associability of the features or elements unique to each.     

Exposure to Spatial Cues Facilitates Visual Discrimination but Not Spatial Guidance

The performance in a radial-arm maze of two groups of rats with restricted access to extra-maze visual cues was studied. One group received extensive exposure to the visual environment of the maze, whereas the second group was never exposed to the environment, aside from their experience in the maze itself. Spatial exposure resulted in a slightly improved ability to discriminate between previously visited and unvisited spatial locations, which can be explained on the basis of general perceptual learning processes. However, there was no evidence that spatial exposure resulted in control of choices by the spatial relations among the maze locations. These results are discussed in terms of theories of spatial learning which appeal to perceptual learning and cognitive mapping.     

Chronic stress leaves novelty-seeking behavior intact while impairing spatial recognition memory in the Y-maze

This experiment examined whether chronic stress disrupts novelty-seeking behavior under conditions that impair spatial memory. Rats were restrained for 6 h per day for 21 days, then tested in either a traditional spatial recognition Y-maze that requires extra-maze spatial cues to navigate or a version with salient intra-maze cues in addition to the extra-maze spatial cues. As previously shown, chronic restraint stress impaired performance on the spatial version of the Y-maze. However, chronically stressed rats performed well in the intra-maze cue version. The results indicate that the deficits in Y-maze performance following chronic stress are not attributed to neophobia, but likely reflect neurochemical and/or neurobiological changes underlying spatial memory ability.     

Acetylcholine release in the hippocampus and striatum during place and response training

These experiments examined the release of acetylcholine in the hippocampus and striatum when rats were trained, within single sessions, on place or response versions of food-rewarded mazes. Microdialysis samples of extra-cellular fluid were collected from the hippocampus and striatum at 5-min increments before, during, and after training. These samples were later analyzed for ACh content using HPLC methods. In Experiment 1, ACh release in both the hippocampus and striatum increased during training on both the place and response tasks. The magnitude of increase of training-related ACh release in the striatum was greater in rats trained on the response task than in rats trained on the place task, while the magnitude of ACh release in the hippocampus was comparable in the two tasks. Experiment 2 tested the possibility that the hippocampus was engaged and participated in learning the response task, as well as the place task, because of the availability of extra-maze cues. Rats were trained on a response version of a maze under either cue-rich or cue-poor conditions. The findings indicate that ACh release in the hippocampus increased similarly under both cue conditions, but declined during training on the cue-poor condition, when spatial processing by the hippocampus would not be suitable for solving the maze. In addition, high baseline levels of ACh release in the hippocampus predicted rapid learning in the cue-rich condition and slow learning in the cue-poor condition. These findings suggest that ACh release in the hippocampus augments response learning when extra-maze cues can be used to solve the maze but impairs response learning when extra-maze cues are not available for use in solving the maze.     

Simplifying environmental cues in a Morris-type water maze improves place learning in old NMRI mice.

Old virgin female NMRI mice aged 17 months were compared with mice aged 3 months for their spatial learning abilities in two versions of the Morris water maze. The first one was a simplified version with a salient configuration of cues comparable to a black/white discrimination and the second one was the classical version of the Morris test with many distal cues surrounding the maze. In the simplified version, old mice presented a slower rate of acquisition and a transient poorer retention compared to young mice. However, old mice achieved a final level of performance statistically comparable to their young counterparts as assessed by latencies to escape onto the concealed platform and by the spatial bias measured in probe trials at intervals during testing. When subsequently subjected to classical Morris maze learning, the same old animals showed marked learning deficits and were persistently impaired in their latencies to escape onto the platform. They presented no spatial bias for the location of the platform in the different probe trials. When the goal was cued at the end of the experiment, the performances of old mice rapidly improved, showing that motivation, motor disabilities, or fatigue and ability to use proximal cues cannot explain the place learning deficit. Our results were discussed in terms of cognitive versus sensory/perceptual disabilities in aged rats and mice.     

Caudate-putamen lesions in the rat may impair or potentiate maze learning depending upon availability of stimulus cues and relevance of response cues

In Experiment 1 rats were required to learn a Y-maze in which reward was made available after a given response (e.g. a left turn) regardless of which arm was used as the start-box. Subjects with lesions of the caudate-putamen showed a deficit on this response-learning task compared with control subjects (unoperated animals and rats having lesions of the posterior cortex). In Experiment 2 rats with caudate-putamen lesions were unimpaired when the direction of the turn required to reach the correct goal-box (identified by means of a salient visual intra-maze cue) varied from trial to trial. In the absence of salient intramaze cues, but with enriched room (extra-maze) cues, the rats with caudate-putamen lesions were superior to controls on this task. It is argued that caudate-putamen lesions disrupt a mechanism responsible for processing information about responses, but that the other (spatial) mechanisms responsible for maze-learning remain intact and that caudate-putamen lesions may enhance performance on spatial tasks for which information about responses is irrelevant.     

Superior discrimination between similar stimuli after simultaneous exposure

Human participants received unsupervised exposure to difficult-to-discriminate chequerboard stimuli (e.g., AX and BX), before learning a discrimination between them. Experiment 1 demonstrated that prior exposure enhanced later discrimination and that intermixed exposure (AX, BX, AX, BX ...) resulted in better subsequent discrimination than did blocked exposure (CY, CY ... DY, DY ...). Experiment 2 showed that simultaneous exposure to two similar stimuli (AX-BX, BX-AX ...) facilitated the later acquisition of a successive discrimination, more than successive exposure (AX-AX, BX-BX ...). These results parallel those observed by Mundy, Honey, and Dwyer (2007) who used pictures of human faces as stimuli and establish the generality of the fact that simultaneous exposure produces a particularly marked perceptual learning effect.     

Effects of Retention Interval on Latent Inhibition and Perceptual Learning

Repeated, non-reinforced preexposure to a context slowed development of conditioned freezing to that context when it was subsequently paired with footshock (latent inhibition) and enhanced discriminability of that context from a similar context (perceptual learning) whether assessed by a generalization test or by explicit discrimination training. Latent inhibition was eliminated by a delay between conditioning sessions and test (Experiments 1a and 1b) and reduced by a delay between preexposure and conditioning (Experiment 2). However, perceptual learning was unaffected by either of these intervals (Experimnets 1b and 2). These results are discussed in terms their impact on theories that have latent inhibition as a possible mechanism of perceptual learning, and on theories of latent inhibition that consider the retardation of conditioned responding to be the result of an acquisition failure.     

Calcium channel antagonists enhance retention of passive avoidance and maze learning in mice

Although a number of studies have shown that treatment with calcium channel antagonists (CCAs) can ameliorate impairments in learning and memory in aged animals, evidence for a general nootropic effect of CCAs in neurologically normal young adult animals is ambiguous. This study attempts to resolve some of this ambiguity by comparing the effects of several CCAs on retention of passive avoidance learning and acquisition and retention of appetitively motivated spatial discrimination learning in young adult mice. Animals were trained in a step through passive avoidance apparatus and, immediately after training, injected subcutaneously with different doses of nimodipine, nifedipine, amlodipine, flunarazine, diltiazem, or verapamil. Retention was tested 24 h after training. In the maze-learning task mice were treated with the same doses of the aforementioned CCAs immediately after a brief training session in a linear maze and retention was tested 24 h after training. The most effective dose of each agent in the maze-retention experiment was administered to additional groups of animals 1 h prior to training to determine the effects of CCAs on acquisition processes. The effects of central administration of CCAs were examined by intracerebroventricular injection of different doses of amlodipine immediately after passive avoidance training. Results showed (1) all peripherally administered drugs except verapamil facilitated retention of passive avoidance training in a dose-dependent manner, (2) all drugs dose dependently facilitated retention of linear maze learning, (3) all doses of the drugs (except verapamil) which facilitated maze retention also facilitated maze learning, and (4) central administration of the dihydropyridine amlodipine produced a dose-dependent facilitation of the retention of passive avoidance learning. These data indicate that drugs which block calcium channels can enhance retention of two different types of learning in mice.     

Children's understanding of death as the cessation of agency: a test using sleep versus death

An important problem faced by children is discriminating between entities capable of goal-directed action, i.e. intentional agents, and non-agents. In the case of discriminating between living and dead animals, including humans, this problem is particularly difficult, because of the large number of perceptual cues that living and dead animals share. However, there are potential costs of failing to discriminate between living and dead animals, including unnecessary vigilance and lost opportunities from failing to realize that an animal, such as an animal killed for food, is dead. This might have led to the evolution of mechanisms specifically for distinguishing between living and dead animals in terms of their ability to act. Here we test this hypothesis by examining patterns of inferences about sleeping and dead organisms by Shuar and German children between 3 and 5-years old. The results show that by age 4, causal cues to death block agency attributions to animals and people, whereas cues to sleep do not. The developmental trajectory of this pattern of inferences is identical across cultures, consistent with the hypothesis of a living/dead discrimination mechanism as a reliably developing part of core cognitive architecture.     

Superior discrimination between similar stimuli after simultaneous exposure

Human participants received unsupervised exposure to difficult-to-discriminate chequerboard stimuli (e.g., AX and BX), before learning a discrimination between them. Experiment 1 demonstrated that prior exposure enhanced later discrimination and that intermixed exposure (AX, BX, AX, BX …) resulted in better subsequent discrimination than did blocked exposure (CY, CY … DY, DY …). Experiment 2 showed that simultaneous exposure to two similar stimuli (AX–BX, BX–AX …) facilitated the later acquisition of a successive discrimination, more than successive exposure (AX–AX, BX–BX …). These results parallel those observed by Mundy, Honey, and Dwyer (2007) who used pictures of human faces as stimuli and establish the generality of the fact that simultaneous exposure produces a particularly marked perceptual learning effect.     

The role of stimulus comparison in human perceptual learning: effects of distractor placement

Within-subjects procedures were used to assess the influence of stimulus comparison on perceptual learning in humans. In Experiment 1, participants received intermixed (A, A', A, A',…) or blocked (B, B,…, B', B',…) exposure to pairs of similar female faces. In a subsequent same/different discrimination task, participants were more accurate when the test involved A and A' than when it involved B and B' (or novel faces: C and C'). This perceptual learning effect was reduced by placing a visual distractor (*: either another face or a checkerboard) between successive presentations of the faces during the exposure stage (e.g., A - * - A'). The attenuation of the intermixed versus blocked difference was particularly marked when faces were used as the distractor. In Experiment 2, this reduction in perceptual learning was more marked when * was positioned between the pairs of intermixed faces (i.e., A - * - A') than when it preceded and succeeded those faces (i.e., * - A - A' - *). These results provide the first direct evidence that the opportunity to compare stimuli plays a causal role in supporting perceptual learning. They also support the specific view that perceptual learning reflects an interaction between a short-term habituation process, that ordinarily biases processing away from the frequently presented common elements and toward their less frequently presented unique elements, and a long-term representational process that reflects this bias.     

Recall of the goal box in latent learning and latent discrimination

Two experiments showed that in a runway rats recalled incidental features of the goal box which they had previously found at the end of it. They were shown to modify their running speed according to the current associated reward value of the goal box. Retrieval cues for features of the goal box were intra- and extra-maze cues, not the response of running. Two goal boxes differing only in colour could be discriminatively recalled by virtue of their association with two discriminable runways. These experiments disconfound recall of the goal box from place learning as factors in latent learning.     

Memory for, and salience of, the unique features of similar stimuli in perceptual learning

In two experiments, participants received exposure to complex checkerboards (e.g., AX and BX) that consisted of small distinctive features (A and B) superimposed on a larger common background (X). Subsequent discrimination between AX and BX, assessed by a same-different task, was facilitated when the stimuli were presented on alternate trials in preexposure--a perceptual learning effect (Experiment 1). The hypothesis that this form of exposure results in more accurate representations of the unique features was supported in Experiment 1, which showed that participants were well able to match the color of the feature with its shape. Experiment 2 showed that exposure to A and B in isolation, intermixed with presentations of AX and BX, enhanced the perceptual learning effect, which confirmed that the better encoding of the unique features during intermixed preexposure is a direct cause of the enhanced discrimination observed following preexposure on this schedule.     

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.