What makes a landmark effective in adolescent and adult rats? Sex and age differences in a navigation task.

In three experiments, rats of different ages were trained in a circular pool to find a hidden platform whose location was defined in terms of a single landmark, a cylinder outside the pool. Following training, two main components of the landmark, its shape and pattern, were tested individually. Experiment 1 was performed by adolescent and adult rats (Exp. 1a, males; Exp. 1b, females). Adult rats always learned faster than the adolescent animals. On test trials, interesting tendencies were found-mainly, one favoring males on the shape test trial, and another favoring females on the pattern test trial. Experiment 2 was conducted only with adolescent rats, and these males and females did not differ when learning the task. However, on test trials the males learned more about the landmark shape component than about the landmark pattern component, while the females learned equally about the two components of the landmark. Finally, Experiment 3 was conducted only with adult rats, and again the males and females did not differ when learning the task. However, on test trials the males learned equally about the two components of the landmark (shape and pattern), but the females learned more about the landmark pattern component than about the landmark shape component. This set of experiments supports the claim that male and female rats can learn rather different things about a landmark that signals the location of the platform, with age being a critical variable.

What makes a landmark effective? Sex differences in a navigation task.

In Experiment 1, two groups of female rats were trained in a triangular pool to find a hidden platform whose location was defined in terms of a single a landmark, a cylinder outside the pool. For one group, the landmark had only a single pattern (i.e., it looked the same when approached from any direction), while for the other, the landmark contained four different patterns (i.e., it looked different when approached from different directions). The first group learned to swim to the platform more rapidly than the second. Experiment 2 confirmed this difference when female rats were trained in a circular pool but found that male rats learned equally rapidly (and as rapidly as females trained with the single-pattern landmark) with both landmarks. This second finding was confirmed in Experiment 3. Finally, in Experiment 4a and 4b, male and female rats were trained either with the same, single-pattern landmark on all trials or with a different landmark each day. Males learned equally rapidly (and as rapidly as females trained with the unchanged landmark) whether the landmark changed or not. We conclude that male and female rats learn rather different things about the landmark that signals the location of the platform.

Associations and propositions: the case for a dual-process account of learning in humans.

We review evidence that supports the conclusion that people can and do learn in two distinct ways - one associative, the other propositional. No one disputes that we solve problems by testing hypotheses and inducing underlying rules, so the issue amounts to deciding whether there is evidence that we (and other animals) also rely on a simpler, associative system, that detects the frequency of occurrence of different events in our environment and the contingencies between them. There is neuroscientific evidence that associative learning occurs in at least some animals (e.g., Aplysia californica), so it must be the case that associative learning has evolved. Since both associative and propositional theories can in principle account for many instances of successful learning, the problem is then to show that there are at least some cases where the two classes of theory predict different outcomes. We offer a demonstration of cue competition effects in humans under incidental conditions as evidence against the argument that all such effects are based on cognitive inference. The latter supposition would imply that if the necessary information is unavailable to inference then no cue competition should occur. We then discuss the case of unblocking by reinforcer omission, where associative theory predicts an irrational solution to the problem, and consider the phenomenon of the Perruchet effect, in which conscious expectancy and conditioned response dissociate. Further discussion makes use of evidence that people will sometimes provide one solution to a problem when it is presented to them in summary form, and another when they are presented in rapid succession with trial-by trial information. We also demonstrate that people trained on a discrimination may show a peak shift (predicted by associative theory), but given the time and opportunity to detect the relationships between S+ and S-, show rule-based behavior instead. Finally, we conclude by presenting evidence that research on individual differences suggests that variation in intelligence and explicit problem solving ability are quite unrelated to variation in implicit (associative) learning, and briefly consider the computational implications of our argument, by asking how both associative and propositional processes can be accommodated within a single framework for cognition.

Landmark vs. geometry learning: explaining female rats' selective preference for a landmark / Aprendizaje basado en un punto de referencia vs.aprendizaje de la geometría: explicando la preferencia selectiva de las ratas hembra por un punto de referencia

Rats were trained in a triangular-shaped pool to find a hidden platform, whose location was defined in terms of two sources of information, a landmark outside the pool and a particular corner of the pool. Subsequent test trials without the platform pitted these two sources of information against one another. In Experiment 1 this test revealed a clear, although selective, sex difference. As in previous experiments, females spent more time in an area of the pool that corresponded to the landmark, but here only when it was a cone but not when it was a pyramid. Males, on the other hand, always spent more time in the distinctive corner of the pool. Experiments 2 and 3 were only with female rats. In Experiment 2 two identical shaped cylinders were used as landmark cues (one plain white and the other vertically patterned with four different patterns). The results of the preference test revealed that only the females trained and tested with the plain cylinder spent more time in the area of the pool that corresponded to the landmark than in the distinctive corner of the pool. Finally, Experiment 3 replicated the results of Experiment 2 while eliminating an alternative explanation in terms of differential contrast between the two cylinders and the black curtain (AU)

Se entrenó a unas ratas en una piscina con forma triangular a que encontrasen una plataforma oculta, cuya ubicación estaba definida en base a dos fuentes de información, un punto de referencia y una parte de la piscina con una forma distintiva. Ensayos de prueba posteriores, sin la plataforma, enfrentaron la forma y el punto de referencia. En el Experimento 1 esta prueba reveló una diferencia de sexo clara, aunque selectiva. Como en experimentos anteriores, las hembras pasaron más tiempo en el área de la piscina que se correspondía con el punto de referencia, aunque sólo cuando este era un cono no cuando era una pirámide. Por otro lado, los machos siempre pasaron más tiempo en el área de la piscina que se correspondía con la forma distintiva. Los Experimentos 2 y 3 se llevaron a cabo sólo con ratas hembra. En el Experimento 2 se emplearon como puntos de referencia dos formas cilíndricas idénticas (una de color blanco y la otra verticalmente dividida en cuatro segmentos con trama diferente). Los resultados de las pruebas de preferencia revelaron que solamente las hembras entrenadas y puestas a prueba con el cilindro blanco pasaron más tiempo en el área de la piscina que se correspondía con el punto de referencia que en el área de la piscina que se correspondía con la forma distintiva. Por último, el Experimento 3 replicó los resultados de los Experimentos 1 y 2 eliminando una explicación alternativa basada en el contraste diferente de los dos cilindros respecto a las cortinas negras (AU)

Do hormonal changes that appear at the onset of puberty determine the strategies used by female rats when solving a navigation task?

The present set of experiments evaluated the possibility that the hormonal changes that appear at the onset of puberty might influence the strategies used by female rats to solve a spatial navigation task. In each experiment, rats were trained in a triangular shaped pool to find a hidden platform which maintained a constant relationship with two sources of information, one individual landmark and one corner of the pool with a distinctive geometry. Then, three test trials were conducted without the platform in counterbalanced order. In one, both the geometry and the landmark were simultaneously presented, although in different spatial positions, in order to measure the rats' preferences. In the remaining test trials what the rats had learned about the two sources of information was measured by presenting them individually. Experiment 1, with 60-day old rats, revealed a clear sex difference, thus replicating a previous finding (Rodríguez et al., 2010): females spent more time in an area of the pool that corresponded to the landmark, whereas males spent more time in the distinctive corner of the pool even though the remaining tests revealed that both sexes had learned about the two sources of information. In Experiment 2, 30-day old female rats, unlike adults, preferred to solve the task using the geometry information rather than the landmark (although juvenile males behaved in exactly the same way as adults). Experiment 3 directly compared the performance of 90- and 30-day old females and found that while the adult females preferred to solve the task using the landmark, the reverse was true in juvenile females. Experiment 4 compared ovariectomized and sham operated females and found that while sham operated females preferred to solve the task using the landmark, the reverse was true in ovariectomized females. Finally, Experiment 5 directly compared adult males and females, juvenile males and females, and ovariectomized females and found that adult males, juvenile males and females, and ovariectomized females did not differ among them in their preferred cue, but they all differed from adult females.

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.

Overshadowing and blocking between landmark learning and shape learning: the importance of sex differences.

Rats were trained in a triangular-shaped pool to find a hidden platform that maintained a constant relationship with two sources of information, an individual landmark and one part of the pool with a distinctive shape. In Experiment 1, shape learning overshadowed landmark learning but landmark learning did not overshadow shape learning in males, while landmark learning overshadowed shape learning but shape learning did not overshadow landmark learning in females. In Experiment 2, rats were pretrained either with the single landmark relevant or with the shape relevant, in the absence of the alternative cue. Final test trials, without the platform, revealed reciprocal blocking only in females; in males, shape learning blocked landmark learning, but not viceversa (Experiment 2a). In Experiment 2b, male rats received a longer pretraining with the single landmark relevant, and now landmark learning blocked shape learning. The results thus confirm the claim that males and females partially use different types of spatial information when solving spatial tasks. These results also agree with the suggestion that shape learning interacts with landmark learning in much the same way as does learning about any pair of stimuli in a Pavlovian conditioning experiment.

Sex differences in the strategies used by rats to solve a navigation task.

Rats were trained in a triangular-shaped pool to find a hidden platform, whose location was defined in terms of two sources of information, a landmark outside the pool and a particular corner of the pool. Subsequent test trials without the platform pitted these two sources of information against one another. This test revealed a clear sex difference. Females spent more time in an area of the pool that corresponded to the landmark, whereas males spent more time in the distinctive corner of the pool even though further tests revealed that both sexes had learned about the two sources of information by presenting cues individually. The results agree with the claim that males and females use different types of information in spatial navigation.

Varieties of perceptual learning.

Although most studies of perceptual learning in human participants have concentrated on the changes in perception assumed to be occurring, studies of nonhuman animals necessarily measure discrimination learning and generalization and remain agnostic on the question of whether changes in behavior reflect changes in perception. On the other hand, animal studies do make it easier to draw a distinction between supervised and unsupervised learning. Differential reinforcement will surely teach animals to attend to some features of a stimulus array rather than to others. But it is an open question as to whether such changes in attention underlie the enhanced discrimination seen after unreinforced exposure to such an array. I argue that most instances of unsupervised perceptual learning observed in animals (and at least some in human animals) are better explained by appeal to well-established principles and phenomena of associative learning theory: excitatory and inhibitory associations between stimulus elements, latent inhibition, and habituation.

Influence of massed and distributed context preexposure on contextual fear and Egr-1 expression in the basolateral amygdala.

Preexposure to the conditioning context can influence the expression of context-conditioned fear. We used behavioral and early growth response gene (egr-1) assays in rats to study the effects of massed and distributed context preexposure on context-conditioned fear. The results demonstrated that massed context preexposure impaired acquisition of contextual fear, an effect here referred to as delayed shock deficit. Spaced context preexposure produced similar inhibitory effects. Significantly, the introduction of a brief change of context prior to conditioning completely reversed the deficit induced by massed, but not by distributed, context preexposure. This reversibility was inversely related to the duration of the context shift. The acquisition of context-conditioned fear was associated with enhanced Egr-1 expression in the basolateral amygdala (BLA). No such increase was evident in animals undergoing distributed context preexposure or in those experiencing massed preexposure without change of context. Remarkably, a brief change of context prior to conditioning not only facilitated learning following massed preexposure but also elicited a significant elevation of Egr-1 protein levels in the BLA. The findings shown demonstrated that the inhibitory effects of massed and distributed context preexposure on conditioning could be dissociable both behaviorally and physiologically. We suggest that the delayed shock deficit associated with massed preexposure derives from perceptual fade-out or inattention and its reversal by a brief change of context from attentional recovery.

Spatial integration with rats.

Rats were trained to find the hidden platform in a Morris pool, whose location was defined by reference to a small number of landmarks around the circumference of the pool. In each of three experiments, an experimental group was trained on alternate trials with two different subsets of three of the available landmarks, with the two subsets sharing one landmark in common. When tested with landmarks drawn from both of their training configurations, but without the landmark common to the two sets, they had no difficulty in locating the platform. In Experiment 1, they performed at least as well as a group trained with all the available landmarks present on every trial. In Experiment 2, they performed significantly better than a group trained with two different subsets of landmarks that shared no one landmark in common.

Competition between landmarks in spatial learning: the role of proximity to the goal.

In two experiments, rats were trained to find a hidden platform in a Morris pool in the presence of two landmarks. Landmark B was present on all training trials, on half the trials accompanied by landmark A, on the remainder by landmark C. For rats in Group Bn, B was near the location of the platform; for those in Group Bf, B was far from the platform. Group Bn performed better than Group Bf on test trials to B alone, but significantly worse on test trials to a new configuration formed by A and C. Thus, the spatial proximity of B to the platform affected not only how well it could be used to locate the platform, but also its ability to prevent learning about other landmarks.

The easy to hard effect and perceptual learning in flavor aversion conditioning.

In experiments on the easy to hard effect, pretraining on an easy discrimination results in better performance on a harder version of the discrimination than pretraining on the hard discrimination itself. In addition, some theories posit that unreinforccd preexposure to the easy discrimination should be as effective as differentially reinforced easy pretraining in producing the easy to hard effect. Two experiments on flavor aversion conditioning in rats demonstrated the basic easy to hard effect. However, in neither experiment was easy preexposure more effective than hard preexposure in enhancing learning of the hard discrimination. Indeed, in Experiment 2, rats preexposed to an easy discrimination learned the hard discrimination significantly more slowly than those preexposed to the hard discrimination itself.

Intradimensional and extradimensional shifts in spatial learning.

Animals trained on 2 discriminations learn the 2nd rapidly if the relevant stimuli are from the same dimension as the 1st (an intradimensional or ID shift) but slowly if the relevant stimuli for the 2 problems are from different dimensions (an extradimensional or ED shift). Four experiments examined ID and ED shifts in spatial learning. Rats trained on 2 spatial problems learned the 2nd more rapidly than rats whose 1st problem had been nonspatial. But this difference between ID and ED shifts depended on the spatial relationship between rewarded (S+) and unrewarded (S-) alternatives in the 2 spatial problems. The results imply that rats trained on a spatial discrimination do not learn to attend to all spatial landmarks but only to those that serve to differentiate S+ and S-.

Associative learning and elemental representation: II. Generalization and discrimination.

This paper follows on from an earlier companion paper (McLaren & Mackintosh, 2000), in which we further developed the elemental associative theory put forward in McLaren, Kaye, and Mackintosh (1989). Here, we begin by explicating the idea that stimuli can be represented as patterns of activation distributed across a set of units and that different stimuli activate partially overlapping sets (the degree of overlap being proportional to the similarity of the stimuli). A consequence of this view is that the overall level of activity of some of the units representing a stimulus may be dependent on the nature of the other stimuli present at the same time. This allows an elemental analysis in which provision for the representation of configurations of stimuli is made. A selective review of studies of generalization and discrimination learning, including peak shift, transfer along a continuum, configural discrimination, and summation, suggests that the principles embodied in this class of theory deserve careful consideration and will form part of any successful model of associative learning in humans or animals. There are some phenomena that require an elemental/associative explanation.

Alternating exposure to two compound flavors creates inhibitory associations between their unique features.

Rats were exposed to two compound solutions, saline-lemon and sucrose-lemon. In Group ALT, trials with one solution alternated with trials with the other. Group BLK received all trials with one solution before any trials with the other. Previous retardation tests had implied that only alternating exposure would establish sucrose as an inhibitor of saline. To provide a complementary summation test for this inhibition, in Experiment 1, all the animals received pairings of peppermint and saline and were tested for consumption of peppermint-sucrose under sodium depletion. Consumption was increased by sodium depletion only in Group BLK. In Experiment 2, a retardation test was used to show that presentation of saline-lemon before sucrose-lemon on each exposure day would establish sucrose as an inhibitor of saline. Neither exposure to sucrose-lemon before saline-lemon nor alternating exposure to sucrose and saline alone had the same effect. These results provide support for an associative theory of perceptual learning that suggests that exposure to complex stimuli aids later discrimination partially as a result of establishing inhibitory associations between their unique elements.

No preguntas si tienen un mapa cognitivo, sino cómo encuentran su camino / Do not ask whether they have a cognitive map, but how they find their way about

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Evidence for inhibitory associations between the unique elements of two compound flavours.

In each of two experiments, rats were pre-exposed to two flavoured solutions, saline-lemon and sucrose-lemon. For group ALT, trials with one solution alternated with trials with the other. Group BLK received all trials with one solution in a block, before any trials with the other. An associative theory suggests that the alternating, but not the blocked, schedule would establish an inhibitory association between sucrose and saline. To provide a retardation test of this inhibition, some animals in each group were then given a single pairing of saline and sucrose, experienced sodium depletion, and were finally tested for their consumption of sucrose. Sodium depletion increased consumption of sucrose more in group BLK than in group ALT. In groups given no saline-sucrose pairing, sodium depletion had only a small effect on sucrose consumption, which was the same in both groups. After multiple pairings of saline and sucrose, sodium depletion had an equally large effect on sucrose consumption in both ALT and BLK groups. These results imply that alternating pre-exposure to two compound solutions does establish an inhibitory association between their unique elements, and thus provide support for an associative theory of perceptual learning.

Is the context specificity of latent inhibition a sufficient explanation of learned irrelevance?

In three experiments, rats were pre-exposed either to uncorrelated presentations of a light and sucrose pellets (group CS/US) or to equivalent presentations of the light and pellets in separate sessions (control). In Experiment 1, subsequent conditioning to the light proceeded more slowly in group CS/US than in the control group, whether this conditioning was excitatory, with the light signalling the delivery of pellets, or inhibitory, with the light signalling their absence. Bonardi and Hall (1996) have argued that this learned irrelevance effect may be reducible to latent inhibition, which would be stronger in group CS/US because they are both pre-exposed and conditioned to the CS in the presence of traces of previous USs occurring in the same session. This analysis implies that group CS/US should have conditioned more rapidly to the CS than controls on the first trial of each session in Experiment 1, but this did not happen. It also implies that the learned irrelevance effect should be reversed if conditioning trials are given at a rate of one per day. Experiments 2 and 3 found no support for this prediction. We conclude that learned irrelevance effects cannot always be reduced to latent inhibition.