Revisiting places passed: Sensitization of exploratory activity in rats with hippocampal lesions

We examined the involvement of the hippocampus in short-term changes in exploratory behaviour in an open field (Experiment 1) and experimental contexts (Experiment 2). In Experiment 1, rats with excitotoxic lesions of the hippocampus were more likely to revisit recently visited zones within the open field than were control rats. Similarly, in Experiment 2 rats with hippocampal lesions showed greater exploration of a context that they had recently explored than a context that they had less recently explored. This short-term sensitization effect was not evident in control rats. These findings are consistent with the suggestion that the recent presentation of a stimulus has two opposing effects on behaviour, sensitization, and habituation, and that hippocampal lesions disrupt the short-term process responsible for habituation, but not that responsible for sensitization.     

Remembrance of places you passed: social spatial working memory in rats

Pairs of rats were tested in a radial-arm maze to determine whether the spatial choices made by one rat affect the subsequent spatial choices of the other rat. In a free-choice procedure, rats showed an increased tendency to choose the location that had most recently been chosen by a foraging partner but a decreased tendency to visit locations that the foraging partner had visited earlier. Forced-choice procedures were used to better control the social stimulus and the interactions between the rats. Under some conditions, locations were chosen later in the choice sequence of a subject rat if another rat had been observed choosing that location. Odor and other physical traces of the other rat's visits were ruled out as explanations for this effect. The results demonstrate the existence of working memory for locations visited by a familiar conspecific.     

Visual discrimination learning and transfer in rats with hippocampal lesions

Two experiments were performed on rats with hippocampal brain damage and on a control group with neocortical lesions. In the first experiment the hippocampal group learned a difficult visual discrimination as promptly as the controls, and neither group was subsequently impaired by adding relevant or irrelevant background cues to the original stimuli. In the second experiment the animals learned a simultaneous visual discrimination in which the stimuli differed in both brightness and orientation. The hippocampal group was impaired relative to the controls on acquisition, and showed poorer transfer to stimuli differing only in brightness or orientation. The results are incompatible with the hypothesis which attempts to explain the effects of hippocampal damage by a widespread reduction in sensory gating, but they are consistent with a more restricted version of the same hypothesis.     

Role of interference in retention by rats with hippocampal lesions.

Involvement of the hippocampus in memory was studied in the rat by employing a retention task with and without interpolated activity. Rats with extensive damage to hippocampus were able to relearn a preoperatively acquired single-alternation task with savings and to perform the single alternation with relatively long delays at a level similar to that of control subjects. However, hippocampals were more affected than normals by an interpolated activity that interferes with retention. The finding of normal retention combined with increased susceptibility to interference supports the view that the memory impairment in subjects with damage to hippocampus may be due to an excess of interference among stored information.     

Hippocampal damage and exploratory preferences in rats: memory for objects, places, and contexts

Rats have a natural tendency to spend more time exploring novel objects than familiar objects, and this preference can be used as an index of object recognition. Rats also show an exploratory preference for objects in locations where they have not previously encountered objects (an index of place memory) and for familiar objects in contexts different from those in which the objects were originally encountered (an index of context memory). In this experiment, rats with cytotoxic lesions of the hippocampal formation were tested on all three versions of the novelty-preference paradigm, with a 5-min retention interval between the familiarization and test phases. Rats with sham lesions displayed a novelty preference on all three trial types, whereas the rats with hippocampal lesions displayed a novelty preference on Object trials but did not discriminate between the objects on Place trials or Context trials. The findings indicate that hippocampal damage impairs memory for contextual or spatial aspects of an experience, whereas memory for objects that were part of the same experience are left relatively intact.     

Distinguishing between new and used traces: Differential effects of electroconvulsive shock on memories for places presented and places passed

In a radial maze test of spatial memory, rats enter relatively novel arms while avoiding locations visited a few hours earlier. Certainly, new memories are acquired for arms entered during the retention test. However, the mnemonic consequences of avoiding arms previously entered are not as clearly predicted; old memories might remain unchanged and yet guide behavior, or the use of old memories during a retention test might renew such memories. The possibilities were evaluated in two experiments in which rats performed in a 12-arm radial maze. Each day the arms were randomly sorted into three sets: A, B, C. Each trial began with forced choices of the 4 arms in Set A and ended after 4 hr in an 8-choice test in which the 4 arms not yet visited (Set B) contained food reward. When electroconvulsive shock (ECS) occurred immediately after Set A choices, accuracy during the test was high; when ECS was administered 2 hr after Set A, choices during the test were less accurate. Old memories therefore appear to be more susceptible than new memories to ECS-induced amnesia. In other trials, an extra retention test was given at the mid-point (2 hr) of the retention interval; this 8-choice test consisted of the remaining 4 arms (Set C) and the original 4 arms (Set A). When ECS was administered after the intermediate test, memory for arms in Set A was 2 hr old (but had just been used), while memory for arms in Set C was new (0 hr). The retention test 2 hr later (testing B vs. A or B vs. C) revealed that ECS had an amnestic effect on the recently used memory for arms in Set A but had no effect on the newly acquired memory for arms in Set C. With respect to ECS-induced amnesia, therefore, memories used in a retention test resemble memories that have aged more than memories that have been newly acquired.     

Dissociation of data-based and expectancy-based memory following hippocampal lesions in rats

Sham-operated and nonoperated animals or animals with hippocampal lesions were presented with sets of trials to test both expectancy-based and data-based memory within the same task. During the study phase of each trial the animals were presented with a constant sequence of five arms on an eight-arm radial maze followed by a test phase in which a recognition test requiring a win-stay rule was used. Expectancy-based memory was measured during the study phase of the trials as a pattern of correct or incorrect orienting responses in anticipation of the ensuing doors in the constant sequence. Both groups of animals acquired correct orienting responses at the same rate, emitted the same pattern of correct orienting responses, and made the same number and pattern of intralist and extralist intrusion errors. Data-based memory was measured during the test phase of the trial as correct recognition test performance. During the test phase the animals with hippocampal lesions were impaired relative to controls on both immediate and 24-h recognition tests. These results suggest that the hippocampus might mediate only data-based, but not expectancy-based, memory and imply a possible dissociation between expectancy-based and data-based memory systems.     

Preserved learning about allocentric cues but impaired flexible memory expression in rats with hippocampal lesions

Several studies have shown that slight modifications in the standard reference spatial memory procedure normally used for allocentric learning in the Morris water maze and the radial maze, can overcome the classic deficit in allocentric navigation typically observed in rats with hippocampal damage. In these special paradigms, however, there is only intramaze manipulation of a salient stimulus. The present study was designed to investigate whether extramaze manipulations produce a similar outcome. With this aim a four-arm plus-shaped maze and a reference spatial memory paradigm were used, in which the goal arm was marked in two ways: by a prominent extramaze cue (intermittent light), which maintained a constant relation with the goal, and by the extramaze constellation of stimuli around the maze. Experiment 1 showed that, unlike the standard version of the task, using this special training procedure hippocampally-damaged rats could learn a place response as quickly as control animals; importantly, one day after reaching criterion, lesioned and control subjects performed the task perfectly during a transfer test in which the salient extramaze stimulus used during the acquisition was removed. However, although acquisition deficit was overcomed in these lesioned animals, a profound deficit in retention was detected 15 days later. Experiment 2 suggests that although under our special paradigm hippocampal rats can learn a place response, spatial memory only can be expressed when the requisites of behavioral flexibility are minimal. These findings suggest that, under certain circumstances, extrahippocampal structures are sufficient for building a coherent allocentric representation of space; however, flexible memory expression is dependent, fundamentally, on hippocampal functioning.