How do animals actually solve the T maze?

Rats were trained on a reinforced, delayed alternation T-maze task in the presence (cue group) or absence (no-cue group) of salient extramaze landmarks. A surprising finding was that the acquisition and memory performance of the 2 groups did not differ. Manipulations of the extramaze landmarks for the cue group suggested that, although landmarks were used to guide behavior, other sources of information were also used normally. The no-cue group was able to perform the task at above-chance levels even when extramaze, intramaze, and inertial sources of orientation were manipulated. These results suggest that memory performance on the T maze does not rely exclusively on the processing of allocentric spatial relationships in the maze environment.

Hippocampal neurons encode information about different types of memory episodes occurring in the same location.

Firing patterns of hippocampal complex-spike neurons were examined for the capacity to encode information important to the memory demands of a task even when the overt behavior and location of the animal are held constant. Neuronal activity was recorded as rats continuously alternated left and right turns from the central stem of a modified T maze. Two-thirds of the cells fired differentially as the rat traversed the common stem on left-turn and right-turn trials, even when potentially confounding variations in running speed, heading, and position on the stem were taken into account. Other cells fired differentially on the two trial types in combination with behavioral and spatial factors or appeared to fire similarly on both trial types. This pattern of results suggests that hippocampal representations encode some of the information necessary for representing specific memory episodes.

Neurotoxic hippocampal lesions have no effect on odor span and little effect on odor recognition memory but produce significant impairments on spatial span, recognition, and alternation.

Recent work has shown that lesions of the hippocampus in monkeys cause deficits in the capacity to remember increasing numbers of objects, colors, and spatial locations (). However, others have observed that hippocampectomized monkeys can show intact memory for a list of objects or locations (). We wished to explore the effects of hippocampal damage on the capacity of memory in the rodent and, to do so, developed novel "span" tasks in which a variable number of odors or locations had to be remembered. In the odor span task (experiment 1), rats were trained on a nonmatching to sample task in which increasing numbers of odors had to be remembered. Half of the trained rats received ibotenic acid lesions of the hippocampus. Postoperatively, hippocampectomized animals did not differ from control animals even when required to remember up to 24 odors. However, when tested on delayed retention of a list of 12 odors, rats with hippocampal lesions were impaired at a long delay. Also, these rats were impaired on a subsequent test of delayed spatial alternation. In a spatial span task (experiment 2), naive rats were trained on a nonmatching to sample task in which a variable number of locations had to be remembered. After this, half of the animals received ibotenic acid lesions. Postoperatively, hippocampectomized animals performed above chance levels when required to remember a single cup location, but were unable to remember more. Subsequent testing on another spatial delayed alternation task suggested that hippocampectomized rats could recognize, but could not inhibit their approach to previously visited locations.

The global record of memory in hippocampal neuronal activity.

In humans the hippocampal region of the brain is crucial for declarative or episodic memory for a broad range of materials. In contrast, there has been controversy over whether the hippocampus mediates a similarly general memory function in other species, or whether it is dedicated to spatial memory processing. Evidence for the spatial view is derived principally from the observations of 'place cells'-hippocampal neurons that fire whenever the animal is in a particular location in its environment, or when it perceives a specific stimulus or performs a specific behaviour in a particular place. We trained rats to perform the same recognition memory task in several distinct locations in a rich spatial environment and found that the activity of many hippocampal neurons was related consistently to perceptual, behavioural or cognitive events, regardless of the location where these events occurred. These results indicate that nonspatial events are fundamental elements of hippocampal representation, and support the view that, across species, the hippocampus has a broad role in information processing associated with memory.

Cue control and head direction cells.

Previous research has shown that head direction (HD) cells in both the anterior dorsal thalamus (ADN) and the postsubiculum (PoS) in rats discharge in relation to familiar, visual landmarks in the environment. This study assessed whether PoS and ADN HD cells would be similarly responsive to nonvisual or unfamiliar environmental cues. After visual input was eliminated by blindfolding the rats, HD cells maintained direction-specific discharge, but their preferred firing directions became less stable. In addition, rotations of the behavioral apparatus indicated that some nonvisual cues (presumably tactile, olfactory, or both) exerted above chance stimulus control over a cell's preferred firing direction. However, a prominent auditory cue was not effective in exerting stimulus control over a cell's preferred direction. HD cell activity also was assessed after rotation of a novel visual cue exposed to the rat for 1, 3, or 8 min. An 8-min exposure was enough time for a novel visual cue to gain control over a cell's preferred direction, whereas an exposure of 1 or 3 min led to control in only about half the sessions. These latter results indicate that HD cells rely on a rapid learning mechanism to develop associations with landmark cues.

The effects of disorientation on visual landmark control of head direction cell orientation.

Head direction (HD) and place cells were recorded in rats that had previously exhibited significant acquisition deficits on a radial arm maze task following disorientation treatment. In this study we determined whether this behavioral impairment was associated with a lack of landmark stimulus control over the preferred orientations of HD and place cells. Neurons were recorded as animals retrieved food pellets in a cylindrical apparatus containing a single cue card. Some of these HD cells were also recorded while animals explored an eight-arm radial maze in a similar cue-controlled environment. The stimulus control of the landmarks in each environment was assessed by rotating the landmark and examining the subsequent preferred orientations of HD and place cells. Animals underwent disorientation treatment before and after each recording session. Despite this disorientation, rotation of the cue card in the cylindrical apparatus resulted in a corresponding shift in the preferred orientations of HD and place cells in 13 of 15 and 7 of 7 recording sessions, respectively. On the radial arm maze, rotation of the landmark cue was associated with a corresponding shift in the HD cell's preferred orientation in 7 of 9 sessions. These results suggest that a visual landmark's stimulus control may not require a learned association between that landmark and an animal's stable experience in an environment. Furthermore, instability in the HD cell system is unlikely to account for the impaired performance of the disoriented animals in the radial arm maze. Rather, these impairments may be due to the animal's inability to utilize stable representations of the environment provided by HD and place cells.

Effects of repeated disorientation on the acquisition of spatial tasks in rats: dissociation between the appetitive radial arm maze and aversive water maze.

This study examined the effects of disorientation on the acquisition of different spatial reference memory tasks. In an appetitively motivated radial arm maze task in which 1 arm was consistently baited, rats that were disoriented before each trial were impaired in their ability to acquire the task relative to rats placed in a clear container and not disoriented. However, disoriented rats were able to learn a Morris water maze and a water version of the radial arm maze under similar training conditions, suggesting that the effects of disorientation may interact with the quality or quantity of motivation involved in a given task. These results suggest that appetitive and aversive spatial tasks are dissociable, and that any impairment that is due to disorientation is specific to the appetitive radial arm maze task.

Correlation between head direction cell activity and spatial behavior on a radial arm maze.

This study assessed the activity of head direction (HD) cells during performance of a spatial reference memory task on a radial arm maze. Rats were trained to select a maze arm located in a constant position in relation to a salient extramaze visual landmark. HD cell discharge properties remained relatively stable across task acquisition in most rats. Following acquisition, rotation of the landmark by 90 degrees or 180 degrees usually led to a corresponding shift in the maze arm selected and the HD cell's preferred firing direction. When the cell's preferred direction did not shift, rats usually selected the wrong arm. HD cell activity was not influenced by the rat's approach to the goal, reward consumption, or exit from the reward area. This demonstration of landmark control over behavior and the cell's preferred direction supports the hypothesis that HD cells contribute to an absolute representation of the environment that can be used to guide spatial behavior.

Processing the head direction cell signal: a review and commentary.

Animals require information about their location and directional heading in order to navigate. Directional information is provided by a population of cells in the postsubiculum and the anterior thalamic nuclei that encode a very accurate, continual representation of the animal's directional heading in the horizontal plane, which is independent of the animal's location. Recent studies indicate that this signal 1) arises either in the anterior thalamic nuclei or in structures upstream from it; 2) is not dependent on an intact hippocampus; 3) receives sensory inputs from both idiothetic and landmark systems; and 4) correlates well with the animal's behavior in a spatial reference memory task. Furthermore, HD cells in the anterior thalamic nuclei appear to encode what the animal's directional heading will be about 40 ms in the future, while HD cells in the postsubiculum encode the animal's current directional heading. Both the electrophysiological and anatomical data suggest that the anterior thalamic nuclei and/or the lateral mammillary nuclei may be the sites of convergence for spatial information derived from landmarks and internally-generated cues. Current evidence also indicates that the vestibular system plays a crucial role in the generation of the HD cell signal. However, the notion that the vestibular system is the sole contributor to the signal generator is difficult to reconcile with several findings; these latter findings are better accounted for with a motor efference copy signal.