Performance ofBetta splendens in a radial arm maze

Siamese fighting fish (Betta splendens) were tested in aquatic versions of radial arm mazes. In the first experiment, the fish were trained to find tubifex worms in an eight-arm maze in which the optimal strategy was to choose each arm once without repetition. After initial training, the fish entered approximately 6.63 different arms in eight choices, showing a strong tendency to choose sequences of adjacent arms, moving about the maze in a Stereotypic direction. This algorithmic response pattern was not, however, sufficient to predict the high performance level of the fish. In the second experiment, a delay of .5 or 5 min was interposed between the fourth and fifth choices. Similar Stereotypic patterns continued in Experiment 2, but choice accuracy following the longer delay declined to a level not significantly above chance. In the third experiment, different fish were tested in a three-arm maze, reinforced either for returning from the second arm to the arm in which they had most recently been fed (win-stay) or for visiting a third arm (win-shift). The fish were significantly faster at acquiring the win-shift contingency than the win-stay contingency. These results demonstrate that solution of spatial tasks depends on the interaction of appropriate behavioral strategies and cognitive capacities that may have little generality across species.     

The effects of maze-arm length on performance in the radial-arm maze

Rats trained in a 16-arm radial maze with arms half the standard length demonstrated extremely low, but above chance, choice accuracy (Experiment 1). Rats trained in a 12-arm maze with short arms demonstrated a substantially higher degree of adjacent-arm responding than did rats trained in the same maze with long maze arms and, when response stereotypy was disrupted by a forced-choice procedure, the short-arm group chose less accurately than the long-arm group (Experiment 2). In a 16-arm maze with 8 short arms and 8 long arms, there was a strong preference for short arms and no evidence for a difference in the ability to discriminate previously visited arms from unvisited arms as a function of arm length, as measured by a two-alternative forced-choice procedure (Experiment 3). These results are interpreted as indicating that arm length affects a choice criterion, with a relatively lax criterion being applied to shorter arms.     

Response cost and time-place discrimination by rats in maze tasks

Time-place discrimination has been shown reliably in several avian and insect species, but only occasionally in rats and fish. In the present experiments, we explored the effects of response cost on time-place discrimination by rats. In the first experiment, we increased the cost of making a choice and the cost of recovering from a wrong choice in two types of maze, a radial arm and a vertical maze. In the radial arm maze, we found only general place preference, whereas in the vertical maze, we obtained evidence of time-place discrimination. In the second experiment, we found that the proportion of rats showing time-place discrimination increased with the height and, therefore, the response cost of the vertical maze. These results suggest that rats do not automatically store and/or retrieve the time and place of reward events but that response cost is an important trigger for time-place discrimination.     

Social effects on spatial choice in the radial arm maze

Social memory was investigated in the context of a spatial working memory task. Pairs of rats were tested in an eight-arm radial maze. Under most conditions, there was a tendency to choose maze locations that had been visited earlier by the other rat. The possibility that this tendency is produced by common preferences for particular maze locations was ruled out. An opposite tendency to avoid visits to locations that had been visited earlier during the trial by another rat was found only when the maze location contained two pellets (rather than an undepletable supply), the rats’ ability to see each other in the maze was restricted to the central arena, and the maze location had been previously visited by the focal rat. The amount of food available in maze locations did not otherwise modulate social influences on spatial choice. The results indicate that memory for a rat’s own previous choices is combined with memory for the choices made by another rat.     

Rats remember not wisely but too well

Rats were trained in a three-alternative spatial delayed matching-to-sample task in a starburst maze. Samples consisted of rewarded forced choices of one arm, and retention was indicated by rats’ returning to that arm after a 90-sec delay. If a rat made an error on its first choice, it was returned to the start compartment and allowed a second choice. Unlike in previous experiments with this task, all three arms were available during the animals’ second choices. The rats tended to perseverate in their second choices by returning to the arm that they had erroneously visited on their first choice. In Experiment 1, the accuracy of second choices following first-choice errors was below chance during the first block of sessions, when a 90-sec delay intervened between the first choice and the second choice, and at chance during the second block of sessions, when a short (5–6 see) delay intervened between first and second choices. In Experiment 2, long-delay and short-delay sessions were randomly presented to naive subjects. Similar results were obtained. In both experiments, the tendency to repeat the erroneous first choice was greater when long delays separated the two choices than when short delays were used. The results suggest that rats make their first-choice errors because they erroneously encode or remember the location of the sample and that they base their second choices on the same erroneous-memory. The increase in perseveration at long delays implies some kind of rehearsal-like mechanism that slows forgetting of the memory controlling the first choice.     

Foraging for covered and uncovered food on a radial maze

Rats were trained to forage for food on a four-arm radial maze. Each arm of the maze was defined as a patch and contained four feeding stations. Each patch contained a total of 20 45-mg food pellets, with the first feeding station in each patch baited with 1 pellet and the remaining stations baited with 1, 5, or 13 pellets. In Experiment 1, one group of rats was tested with feeders open and food readily accessible, and another group was tested with metal covers on the feeders, which necessitated extra time to gain access to food. With open feeders, the rats visited each feeder in a patch in the order in which they encountered the feeders, from the center of the maze to the end of the arm. The rats in the group with the covered feeders often visited the feeders containing 5 or 13 pellets first and the feeders containing 1 pellet last. In Experiment 2, it was found that the rats switched readily between these two foraging strategies when tested with covered and open feeders on alternate sessions. The extra time and effort required to uncover food appeared to produce selective foraging in rats.     

Retroactive inhibition in rat spatial memory

Two experiments were carried out in which rats first were given four forced choices on an eight-arm radial maze, then were given interpolated maze experiences, and finally were given a free choice retention test on the first maze. In Experiment 1, interpolated experiences consisted of forced choices made on one, two, or three other mazes, each placed in a different room. Retroactive inhibition (RI) was not found with one and two interpolated mazes but was found with three interpolated mazes. In Experiments 2a and 2b, an attempt was made to produce RI within a single context by using two mazes placed side by side or on top of one another and by using interpolated forced choices that were different, random, or the same with respect to forced choices onMaze 1. These conditions failed to yield any evidence of RI. In Experiment 2c, forced choices were followed by interpolated direct placements on the same maze on different, random, or the same maze arms, and retention tests revealed RI under these conditions. It was concluded that rats encode memories of specific places visited in space and that RI will arise only if (1) memory is greatly overloaded with interpolated information or (2) an interpolated visit is made to exactly that position in space to which an animal must travel in order to achieve a correct choice on the retention test.     

Intratrial proactive interference in rats’ serial alternation performance in the radial maze

Rats acquired a serial alternation task in an eight-arm radial maze that was partitioned into four pairs of arms. Each pair was associated with a different distal stimulus. Rats were initially forced to the left or right arm in each pair (the study segment) before being exposed to both arms in each pair (the free-choice or test segment). Only the previously blocked arm of each pair remained baited. Following initial training, proactive interference (PI) was induced by presenting rats with a forced-choice (prestudy) segment containing arm positions opposite those in the subsequent study segment. Such trials generated poorer free-choice accuracy than did trials without a prestudy segment. Forcing rats to both arms in the pair in a prestudy segment produced only transient PI. A slight improvement in rats’ free-choice performance was obtained by forcing them to the same arm position, but only when the test segment was delayed by 30 min. Increasing the interval between the prestudy and study segments from 2 to 30 min eliminated PI, but only when free-choice testing was delayed by 2 min rather than by 30 min. These results suggest that intratrial PI in this preparation was primarily due to confusion about which arm position in each pair had been visited during the last forced-choice segment.     

The influence of spatial irregularity upon radial-maze performance in the rat

In two experiments, we examined how the introduction of vertical or horizontal irregularities in the perfectly regular shape of a radial maze affected rats’ performances. The introduction of various tilts in each arm of an eight-arm radial maze had a slightly positive effect on accuracy. However, when intra- and extraraaze cues were dissociated by rotating the maze before maze completion, the rata relied preferentially on extramaze cues associated with the horizontal direction of the arms but not with the tilts. On the other hand, the rats showed poor performances when trained on a horizontally distorted maze (uneven angles between the arms instead of repeated 45° angles). The high number of errors was related to the neglect of particular arms, the disorganization of the patrolling sequences, and the tendency to chain the visit of five arms that formed a regular ahape. Other animals, trained in the same maze, displayed similar biases even after a pretraining phase with constrained choices. Results from the horizontally distorted maze confirm and extend data from the spontaneous alternation literature that choice behavior is influenced by rules of movement that favor large angle transitions and regular subdivisions of space. They also stress the relation between performance in the radial maze and spontaneous exploratory and foraging behaviors.     

Maze-arm length affects a choice criterion in the radial-arm maze

Male rats were tested in a 12-arm radial maze with 6 arms that were standard in length and 6 arms that were half the standard length. As previously reported by Brown (1990), revisits to short arms were more likely than revisits to long arms. Two explanations of this effect of mazearm length on choice accuracy were experimentally contrasted. The first attributes the effect to diminished discriminability of visited and unvisited arms when the arms are short. The second attributes the effect to a relatively lax choice criterion being applied to short arms. An analysis of the microstructure of choices, applying the logic of signal detection theory, provided evidence for the latter explanation.     

Retroactive interference in rat radial maze performance: The role of point of delay interpolation and the similarity and amount of interpolated material

The conditions necessary for producing retroactive interference (RI) were examined in a 12-arm radial maze. Rats were first given either three or nine forced choices in a to-be-remembered maze. During a 2-h delay, they received one or two trials in a second 12-arm maze, located either in a different room or the same room as the to-be-remembered maze. During the postdelay memory test, RI from the interference trials was produced only when nine choices had been made in the to-be-remembered maze and two interference trials had been conducted during the delay interval. RI was not found when only three forced choices had to be retained or after a single interference trial. The similarity between the interpolated and to-be-remembered mazes had no effect on choice accuracy. It was concluded that two conditions are required for the production of RI in the radial maze. First, a “large amount” of information should be resident in working memory. Second, a substantial number of interpolated trials or choices must be made during the delay.     

Intramaze cues and “odor trails” fail to direct choice behavior on an elevated maze

The relative importance of intramaze cues and extramaze cues in directing choice behavior on a radial arm maze was examined using a discrimination procedure which selectively rewarded rats for following only one set of cues. Rats in the intramaze group obtained food from a food cup on the end of each arm. Rats in the extramaze group obtained food from a food cup on a small platform just beyond the end of each arm. All rats were first shaped to perform correctly with the maze in a constant position. Then the maze was rotated to a new position after every choice. For rats in the intramaze group, the food moved with the arms, making intramaze cues relevant. For rats in the extramaze group, the food remained on the platforms (in the same position in the room), making extramaze cues relevant. Rats in the extramaze group performed almost perfectly during maze rotation, demonstrating that intramaze cues were not necessary to support accurate choice behavior. Rats in the intramaze group never performed better than chance, demonstrating that intramaze cues (from the rats, the reinforcement, and the apparatus) were not adequate to control choice behavior. The results of the present experiment are compared to those of other experiments describing the influence of “odor trails” or other olfactory stimuli on choice behavior in mazes.     

Rat spatial memory: Resistance to retroactive interference at long retention intervals

An attempt was made to disrupt memory for spatial information by interpolating a task of high similarity to the to-be-remembered task during a long retention interval. Rats were trained in an 8-arm maze in which choosing each arm without repetition was the optional strategy. A 4-h delay was imposed between the 4th and 5th choices. At various times during the retention interval, the rats ran a second identical maze located in another room. No evidence of retroactive interference was observed. In the second experiment, the rat was required to remember the interpolated spatial task during the retention test. This was accomplished by allowing the rat to make four choices in the first maze and then, after a variable period of time, four choices in the second maze. Four hours after exposure to each maze, retention was tested. Choice accuracy on the retention tests was high and equivalent on both mazes. Requiring the rat to remember which arms it had visited in a second maze did not impair memory for the first maze. These results demonstrate that rats can segregate spatial memories established in different contexts with considerable proficiency.     

Foraging on the radial-arm maze: Effects of altering the reward at a target location

Thirsty rats were trained to collect small water rewards from the end of each arm of an eight-arm radial maze. During these training trials and subsequent testing trials, the subjects were allowed to choose a maximum of eight arms. “Preference” for a target maze location was studied by noting when, in the sequence of eight choices, the target was selected. During testing, when one maze location was consistently devoid of water, rats decreased their preference for this arm over trials (Experiment 1). Similarly, rats that learned a saccharin-lithium association demonstrated lower preferences for a maze location that consistently held the conditioned saccharin solution. This was true for animals that received saccharin-lithium conditioning on the maze (Experiment 3A) and for animals conditioned to saccharin in a separate context (Experiment 3B). An increase in preference for a target maze location consistently containing a sweet chocolate milk solution was observed in animals that were water- and food-deprived (Experiment 2). These studies demonstrate that animals will modify their responses toward (preferences for) maze locations that predictably contain an altered reward.     

Stimulus control and function of arm and wall travel by rats on a radial arm floor maze

Hoffman, Timberlake, Leffel, and Gont (1999) concluded that the tactic of effective trail following (in the form of arm and wall travel), rather than distance minimizing, central-place search, or random search, best characterized the locomotion of rats on a radial arm maze placed flat on the floor of an arena (a floor RAM). The present experiments analyzed further the stimulus control and function of arm and wall travel. Experiment 1 showed that arm travel was controlled more by the edge of a maze arm than by its surface. Experiment 2 showed that rats with whiskers clipped on one side traveled along arms less and along walls more than did intact rats. Experiment 3 showed that maze arms increased search effectiveness and decreased suppression of locomotion by bright light and a novel environment. The results support the hypothesis that arm and wall travel are based on mechanisms of trail following, which, in natural settings, contribute to food finding and regulation of social relations and fear.     

Maze patrolling by rats with and without food reward

The effects of food reward on rats’ behavior in radial and Dashiell tunnel mazes were examined in two experiments. In the first, with animals at ad-lib body weights, food reward reduced speed of movement at the food locations, but did not affect the patterns of movement in either maze. Exploratory efficiency in the Dashiell maze was unaffected by food reward, and spontaneous patrolling of the radial maze by the nonrewarded animals was comparable to the behavior, reported by others, of rats running for food reward on elevated eight-arm mazes. In the second experiment, with subjects maintained at 80% of ad-lib body weights, there was some evidence for “winstay” learning: food-rewarded rats in the Dashiell maze were relatively more active near the food locations than were the nonrewarded animals, and more rewarded than nonrewarded rats revisited all food locations in the radial maze. Nonetheless, exploratory efficiency in the Dashiell maze was unaffected by food reward, as was patrolling efficiency in the radial maze, which was again comparable to that of rats on elevated mazes. The similarity in behavior of rewarded and nonrewarded animals in these mazes implies that the major determinant of their behavior, whether or not food reward is provided, is a spontaneous tendency to avoid places recently visited.     

Spatial memory in rats: Resistance to retroactive interference

Treatments that interfere with animals’ short-term retention (e.g., in delayed matching-to-sample) were studied using a spatial memory task. Rats performed in an eight-arm radial maze in which choosing each arm without repetition was the optimal behavior. Performances were interrupted between fourth and fifth choices for a delay of 15 sec to 2 min. A variety of events occurring during the delay interval did not disrupt memories for prior choices (as assessed by the accuracy of postdelay choices). The ineffective treatments included variations in visual and auditory environments, removal from the maze, food consumed during the delay, a distinctive odor added to the maze, or combinations of these manipulations. Additionally, performance on another spatial task (a four-arm maze) during the delay between Choices 4 and 5 did not interfere with performance in the eight-arm maze. These findings suggest that rats’ memories for spatial locations are immune to retroactive interference, at least within the range of conditions reported, and that the rat can successfully segregate memories for spatial locations established in different contexts.     

Central-place foraging by rats on the radial maze: The effects of patch size,food distribution,and travel time

Rats foraged on a four-arm radial maze with one, two, three, and four food items (0.65.g pieces of cheese) placed on different arms (patches) of the maze. In two experiments, the hypothesis was tested that rats should carry food to the center of the maze more often when a patch contains one food item than when it contains multiple food items. Support for this prediction was found when the tendency to carry initial items encountered in patches was compared among the different sized patches. However, a further observation failed to support the hypothesis: Food carrying declined from first to last item encountered in multiple-item patches with clustered food items. Experiment 1 revealed that food carrying was reduced when travel time was increased by barriers placed at arm entrances. Both Experiments 1 and 2 indicated that the tendency for rats to carry food to the center of the radial maze increased as the distance of food encountered on an arm increased from the center. In both experiments, some rats dealt with the problem of multiple items by resorting to multiple-item loading, and some rats carried food items from the end of an arm to a point on the arm nearer the center for consumption.     

Testing optimal foraging theory on the radial maze: The role of learning in patch sampling

A four-arm radial maze containing 10 feeders in each arm (patch) was used to study patch sampling in rats. In each of three experiments, rats foraged for 30 sessions. On each session, two randomly chosen patches were baited with food and the remaining two patches were empty. In Experiment 1, the number of baited feeders in baited patches (6) was varied from 1–10 over five groups of subjects. Mean visits to empty patches was an inverse function of 6, as predicted by an optimal foraging model. In Experiments 2 and 3, rats’ ability to discriminate between baited and empty patches was examined when food in baited patches was placed in fixed locations, either in clumps (Experiment 2) or distributed throughout the patch (Experiment 3). Rats in fixed-food-location conditions reliably visited fewer feeders in empty patches than did rats in randomly changing control groups. Examination of within-patch foraging patterns indicated that rats in fixed-food-location groups selectively sampled potentially baited locations and abandoned the patch if food was not found. It is suggested that processes of patch discrimination were responsible for these effects.     

Effects of varying trial distribution,intra- and extramaze cues,and amount of reward on proactive interference in the radial maze

Rats are typically less accurate in their arm selections in the radial maze over successive trials in a session (Roberts & Dale, 1981). In the present study, rats’ choice accuracy declined when such trials were separated by 2-min (massed) but not by 2-h (spaced) intertriai intervals. Changing intramaze visual/tactile arm stimuli (Experiments 1 and 3) or extramaze landmark stimuli (Experiment 4) between trials weakened the massed-trials effect, but changing the number of food pellets per arm, either alone or in conjunction with changes in intramaze cues (Experiments 2 and 3), did not. The rats also tended to avoid the spatial locations of their last four choices on a previous trial during their first four choices on a current trial, and more so with massed than with spaced trials. These findings indicate that intertriai proactive interference (PI) occurred only with massed trials and was weakened by changing intra- and extramaze cues between such trials.