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 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.     

Spatial pattern learning in the radial arm maze

Rats experienced a spatial pattern of baited and unbaited arms in an eight-arm radial maze. The spatial pattern remained constant over trials, but the spatial locations that were baited varied unpredictably. Although there was no evidence of control by the spatial pattern during free choice training trials, the rats’ ability to locate baited arms in forced choice test trials was superior to that of animals in a control condition for which maze arms were not baited in a consistent spatial pattern. This is consistent with the results of experiments showing that spatial choices by rats in a pole box maze are controlled by abstract spatial patterns.     

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.     

How does the ecological foraging behavior of desert kangaroo rats (Dipodomys deserti) relate to their behavior on radial mazes?

Experiment 1 showed that laboratory-reared desert kangaroo rats, like domestic Norway rats, efficiently search for food on a radial arm maze (RAM) by avoiding revisiting arms within a trial. By placing an RAM on the floor so the animals could approach food from any direction, Experiment 2 tested whether efficient search by kangaroo rats was based on tactics of distance minimizing, central-place foraging, trail following, or meandering. In contrast to the dominant trail-following tactic of domestic Norway rats (Hoffman, Timberlake, Leffel, & Gont, 1999), kangaroo rats tended to distance minimize, whether maze arms were present or not. Experiment 3 indicated that kangaroo rats treated a floor configuration of eight food cups as twopatches of four, based on beeline travel between patches and meandering within them. We conclude that similar performance in an elevated RAM by different species can be based on different tactics, and we suggest that a laboratory apparatus can be used to cast light on niche-related mechanisms.     

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.     

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.     

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.     

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.     

Exploring the limits of spatial memory in rats,using very large mazes

In Experiment 1, rats foraged for food in six successive phases with 8, 16, 24, 32, 40, and 48 arms attached in random locations to a large radial maze. The percentage of novel choices appeared to be determined more by spatial proximity than by number of arms. In Experiment 2, rats foraged for food in four successive phases with 8, 16, 24, and 48 arms attached to the maze in spread-out or tight configurations. Performance was poor in the tight configurations regardless of the number of arms. Performance was excellent in the 8-arm spread-out condition but declined as 16 and, then again, 24 arms were added. Thus, spatial separation, not number of locations, was the chief determinant of performance in the first two experiments. In Experiment 3, in successive phases, 8, 16, 24, 32, 40, 48, 16, and 8 food towers were set in a circle on the floor, with the spatial separation between adjacent towers held constant at 33 cm. The percentage of novel choices declined as 8 towers became 16 and did not change again with 24, 32, 40, or 48 towers in place but then increased again as 16 towers became 8. In Experiment 4, in successive phases, 8, 16, 24, and 32 food towers were set in a circle, with the spatial separation between adjacent towers held constant at 66 cm. The percentage of novel choices declined as 8 towers became 16 and again as 16 towers became 24 but did not decline further. These data were discussed in terms of the fundamental problems posed by variations in the number of food locations in the pursuit of the limit of spatial memory in rats.     

How is radial arm maze behavior in rats related to locomotor search tactics?

Norway rats have been shown to depend on short-term spatial memory to find food on a radial arm maze (RAM), but what locomotor search tactics are involved in using this memory effectively? Four experiments distinguished tactics of distance minimizing, central-place search, trail following, thigmotactic search, and random search by using different configurations of a RAM placed flat on the floor of an arena. These search tactics make similar predictions on an elevated RAM but predict different outcomes on a floor RAM because the rats are free to approach the food from any direction. After initial trials dominated by exploration, rats traveled along arms to food, even when the resultant distance was up to three times the minimum distance. With no food present, rats also traveled along arms; with no arms up to present, they traveled along walls to food. It appears that both maze arms and arena walls engage mechanisms related to trail following in rats.     

Effects of runway shift and stay rules on rats’ serial pattern learning in the T-maze

Rats received three-trial series on a T-maze consisting of extended visually distinct left-black and right-striped side runways. During the first phase of training, when allowed to select baited runways within these series, they predominantly alternated their choices. During the second phase, rats received forced-choice serial pattern training of series consisting of two rewarded (R) trials and one nonrewarded (N) trial in two fixed orders, RRN and RNR. In Experiment 1, the rats in the runway shift rule group always received the second R trial when forced down a runway opposite that on the preceding trial in the series and the N trial when forced down the same runway. The rats in the runway stay rule group always received the second R trial when forced down the same runway and the N trial when forced down the opposite runway. In Experiment 2, each rat was conditionally trained with both runway outcome rules as determined by the central alley lighting and the type of food in the side alleys. The rats took longer to reduce their running speed on the N trial within each sequence under the runway stay rule than under the runway shift rule. They also took longer to acquire serial pattern responding for the RNR than for the RRN series only under the runway stay rule condition. When subsequently reexposed to series of free-choice trials on the final phase, rats maintained spontaneous alternating choice patterns under the runway shift rule conditions but either seldom alternated their choices (Experiment 1) or greatly reduced choice alternations (Experiment 2) under the runway stay rule condition. We discussed these effects in terms of rats’ natural foraging strategies and as a factor that interacts with other within- and between-series variables that affect serial pattern behavior.     

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.     

Multiple-pattern learning by rats on an eight-arm radial maze

Rats were trained over a number of sessions on an eight-arm radial maze with eight trials on each session. Each of four arms on the maze contained a different pattern formed by sequences of reward (two pellets) or nonreward (no pellets) over successive trials within sessions. The patterns were single alternation, double alternation, and two patterns in which four rewards or four nonrewards were preceded and followed by two nonrewards or two rewards, respectively. The other four arms on the maze served as control arms and always contained one pellet. It was found that rats tracked all of these patterns when they were required to climb over barriers to enter and leave arms. However, rats showed no ability to extrapolate patterns beyond the training trials. These findings, and a further analysis of arm-choice stereotypy, led to the conclusion that rats tracked by using a trial-number strategy.     

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.     

Rats show preference for delayed rewards on the radial maze

Rats on an eight-arm radial maze chose between four arms on which a small reward could be obtained after a short delay and four arms on which a larger reward could be obtained after a longer delay. Experiments 1 and 2 showed that rats preferred the long-delay, large-reward arms over the short-delay, small-reward arms. This preference was particularly marked when the arms were made into enclosed alleys. Experiment 3 showed that this effect was not produced by a preference for staying in enclosed alleys. We argue that the rats endured longer delays to obtain larger rewards when fear of predation was minimized.     

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.     

Disruption of temporal discrimination and the choose-short effect

The present experiment examined the effects of several disruptors on temporal discrimination. Pigeons responded under a 0-delay symbolic matching-to-sample procedure in which responses to one key color were reinforced following the presentation of four shorter sample durations, and responses to another key color were reinforced following the presentation of four longer sample durations. Steady-state performance was disrupted by presession feeding, intertrial-interval food, visual distraction, and extinction. All disruptors decreased temporal-discrimination accuracy. Analyses of the fitted cumulative normal functions indicated that decreases in accuracy were produced mainly by decreases in overall stimulus control rather than specific effects on timing. In addition, all disruptors selectively decreased accuracy on long-sample trials—a choose-short effect. This effect is interpreted in terms of decreased attention to the samples under disruption. Current theories of the choose-short effect do not appear to easily account for these results.     

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.     

The influence of cue type and configuration upon radial-maze performance in the rat

The influence of cue type and cue configuration on radial-maze performance in rats was examined in two experiments. In the first experiment, it was found that rats provided with both salient intramaze and extramaze cues acquired the task faster than rats given only one set of cues. No difference in acquisition was found between a group trained with intramaze cues alone and a group trained with extramaze cues alone. In a cue-preference test, it was found that groups that had been trained with extramaze cues, intramaze cues, or both sets of cues relied on extra-maze cues to avoid visited arms when given both types of cues concurrently. When all groups were transferred to intramaze-cue-alone trials, only the group that had been originally trained with extramaze cues alone showed any disruption. Also, during the second half of the intramaze-cue-alone trials, the arrangement of these cues was randomly changed on each trial. This disruption in cue configuration did not deleteriously affect performance in any of the three groups; all remained above chance performance, although the performance of the group originally trained with extramaze cues alone was inferior to that of the other two groups. In Experiment 2, groups of rats were trained on daily alternating trials under intramaze-cue-alone and extramaze-cue-alone conditions. For one group, the configuration of intramaze cues was altered randomly on each trial; the other group had intramaze cues always presented in the same configuration over trials. It was found that acquisition was more rapid on intramaze trials in the group given static configurations. Also, acquisition of the extramaze task was faster than the intramaze task in the group given variable intramaze cue configurations. No difference was found between the intramaze and extramaze conditions in the group given static intramaze cue configurations. These data suggest that a static cue configuration may influence radial maze performance, but is not a necessary condition for such performance.