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.     

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.     

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.     

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.     

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.     

In the dark: Spatial choice when access to spatial cues is restricted

The cognitive mechanisms involved in spatial choice when access to visual cues is restricted were examined in three experiments using male rats. A specially constructed radial-arm maze was used, in which extramaze visual cues could not be perceived from the central arena, but could be perceived from the maze arms. Choices were very accurate when the maze was not rotated during each trial, but inaccurate when the maze was rotated. This suggests that intramaze cues were involved in the control of choices. However, the data clearly showed that choices were not simply controlled by intramaze cues. Rather, control by intramaze cues interacted in a more complex manner with representations of the spatial locations of goals. Such spatial representations were involved in the control of choice despite the absence of visual spatial cues at the time choices were made.     

The sensory basis of spatial memory in the rat

Rats were given three stages of training on an eight-arm, elevated radial maze with food reward at the end of each arm. In Stage 1, rats were allowed to choose freely among the arms from the beginning of a trial. In Stage 2, three initial forced choices were followed by a series of free choices. In Stage 3, the central platform of the maze was rotated with the rat on it between the initial forced choices and the free choices. Following testing on these three stages, the animals were divided into four groups and deprived of selected senses. One group was made blind, a second anosmic, a third blind and anosmic, and a fourth was left normal. The same three stages of testing that had been conducted preoperatively then were run again post-operatively. Throughout these tests, the possible use of auditory cues was tested by presenting white noise on alternate trials. Finally, two further tests were carried out, the multiple rotations test and the removal-replacement test. The results indicated that visual cues, but not olfactory or auditory cues, played a critical role in the rat’s ability to avoid previously entered alleys. There was evidence also that rats used internal cues from kinesthetic and/or vestibular receptors when visual cues were absent.     

Spatial configuration and list learning of proximally cued arms by rats in the enclosed four-arm radial maze

In an enclosed four-arm radial maze, after sampling three experimenter-selected baited arms (thestudy segment) and following rotation of the maze, rats had to find the fourth baited arm among all four unblocked arms (test segment). The rats learned this task with two sets of arm cues, objects at arms’ entrances and full arm inserts, each maintained in a fixed configuration. When we changed the configuration of one set of arms to itsmirror image and that of the other set to a moremixed variation by switching opposite and adjacent cued arms, the rats’ accuracy was similarly disrupted (Experiment 1). In Experiment 2, the same rats rapidly recovered their high search accuracy on four new configurations recombined from pairs of adjacent arms and pairs of opposite cued arms from the previous final two configurations. Their test segment search accuracy, however, was again disrupted when these configurations were varied either only over trials’ study segments or only over trials’ test segments. In Experiment 3, however, these rats attained accuracy as high on two sets of cued arms with constantly changing configurations as on two sets with constant configurations. Thus, the rats were able to separately represent four different spatially stable configurations, and then they could learn to represent two of these configurations as lists of spatially irrelevant items. We discuss these findings in terms of association theory and parallel map theory (Jacobs & Schenk, 2003).     

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.     

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.     

Evidence for a shift in the choice criterion of rats in a 12-arm radial maze

Rats were trained in a standard 12-arm radial maze task. Following training, each trial consisted of a sequence of 2, 4, 6, 8, or 10 choices, followed by a 15-min delay, which then was followed by a choice between a single arm and a response manipulandum mounted in the center of the maze. An arm visit was reinforced if the arm had not been visited prior to the delay, whereas a manipulandum response was reinforced if the arm had been visited. It was found that rats are relatively more likely to reject arms by responding to the manipulandum following a delay occurring late in the choice sequence. This indicates that the choice criterion used by rats in the radial maze becomes more strict as the choice sequence progresses. Such a process provides an alternative explanation for some of the data recently reported by Cook, Brown, and Riley (1985).     

The role of extramaze cues in spontaneous alternation in a plus-maze

Performance on various spatial tasks is superior in the presence of extramaze cues that provide guidance in locating the goal. This study was an investigation of the effect of such cues on nonrewarded, continuous spontaneous alternation in a plus-maze, a spatial task with no explicit reinforcement. Rats were tested in five daily sessions. Alternation scores, which reflect how frequently the least recently visited arms are chosen, were calculated for each day. On the first day only, rats in the stimulus-rich condition, with cues added to the curtain that surrounded the maze, alternated more than did rats in the stimulus-poor condition. Over days, the stimulus-rich group showed a significant downward trend in alternation, whereas the stimulus-poor group showed an upward trend, but all the groups always alternated at levels greater than chance. Thus, although performance on this task can be enhanced by the presence of extramaze cues, other types of information support alternation.     

In the dark II: spatial choice when access to extrinsic spatial cues is eliminated

Rats were tested in a specially constructed radial-arm maze that eliminated access to extramaze visual cues and allowed any effects of intramaze cues to be controlled. Despite this, choice accuracy was controlled by the spatial location of previously visited arms. Part of this control was attributed to vestibular or kinesthetic cues. This conclusion was corroborated by the finding that when explicit visual cues were moved from their standard (trained) spatial locations to novel locations, control of spatial choices was completely disrupted. The latter finding indicates that cues intrinsic to the rat (kinesthetic or vestibular information) and cues extrinsic to the rat (visual stimuli) operate in an integrated fashion.     

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.     

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.     

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.     

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.     

A “random-walk” simulation model of multiple-pattern learning in a radial-arm maze

Wathen and Roberts (1994) reported rather surprising results of a radial maze study that on any interpretation requires postulation of previously unsuspected high-level cognitive processes in rats. In each of four arms of the eight-arm radial maze, a different serial pattern unfolded over trials; for example, in one of the arms reward and nonreward alternated over successive trials. On each trial, rats came to track successfully four different patterns simultaneously. The authors suggested that rats tracked the pattern by using some form of trial-number strategy; that is, the trial number indicated which arms contained the better rewards. This strategy could involve a hypothesis, considered unlikely by some, that rats are capable of keeping track of as many as eight successive events—as, for example, by counting. A simulation model that embodies a specific form of the trial-number hypothesis is described here, and the results of the simulation correlate remarkably well with the observed data. In addition, the model makes four separate predictions that are supported by Wathen and Roberts’s data and that seem beyond the scope of other available theories.     

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.     

Spatial localization of a goal: Beacon homing and landmark piloting by rats on a radial maze

We performed six experiments in order to examine the ability of rats to use moving beacons and landmarks as cues to the location of reward on an eight-arm radial maze. In Experiments 1–4, the cues and goals were moved before each trial, and groups in which a single beacon was placed on the rewarded arm, a single landmark indicated that reward was on the arm immediately to the left of a landmark, or two landmarks were placed on each side of the reward arm were compared. The rats rapidly learned to track the reward in the beacon condition, failed to find the reward sooner than chance expectation with a single landmark, and did only slightly better than chance with two landmarks. In Experiments 5 and 6, the rats were trained in five trials per day, with the landmark and goal locations constant over daily rewarded trials, and in two extinction trials that were inserted among the rewarded trials. The rats found the goal arm at substantially better than chance expectancy with both one and two landmarks. Our results, in agreement with data from recent swimming pool experiments (A. D. L. Roberts & Pearce, 1998), show that rats will use the relationship between moving landmarks and a goal in order to find reward.