Further studies of pigeons’ spatial working memory in the open-field task

In Experiments 1 and 2, pigeons’ spatial working memory in an open-field setting was examined under conditions that differed in terms of working-memory load (number of sites visited prior to a retention test) at various delays between initial choices and the retention test. In Experiment 1, pigeons were tested under two conditions of memory load (three or five sites visited prior to the delay) and two delay intervals (15 and 60 min). Accuracy declined as a function of delay but was not affected significantly by memory load. In Experiment 2A, pigeons were tested under three conditions of memory load (two, four, or six sites visited prior to the delay). In separate phases, the delay was 2, 15, and 60 min. Accuracy was not affected by memory load in any of these phases. In Experiment 2B, three conditions of memory load (two, four, or six sites visited prior to the delay) were tested at two delays (2 and 60 min) within a test phase. Accuracy declined with increasing delay, but memory load again had no significant effects. These results are inconsistent with previous suggestions that pigeons’ retention of spatial information may decline as working-memory load is increased. In Experiment 3, cue-manipulation tests confirmed that pigeons’ choice behavior in the open-field task is controlled by memory for previously visitad room locations.     

More evidence of robust spatial associative memory in the pigeon,Columba livia

Willson and Wilkie (1993) developed a novel procedure for assessing pigeons’ memory for the spatial location of food. Only one of four locations (consisting of an illuminated pecking key and grain feeder) provided food each day. Over days, different locations provided food. The pigeons’ tendency to revisit the location that was profitable on the previous day demonstrated memory for food-spatiallocation associations over a period of 24 h, retention longer than previously reported for this species. This basic finding was replicated and extended in three experiments. Experiment 1 demonstrated that location-food discriminations were also remembered well when established with successive rather than concurrent procedures. Experiment 2 demonstrated that pigeons can remember two location-food associations over 24 h. Experiment 3 showed that the discrimination training inherent in this paradigm is important for retention; retention was impaired when only the rewarded location was presented. Overall, this research suggests that cross-species differences in spatial memory performance may be due to quantitative rather than qualitative differences in the memory system underlying performance.     

No evidence for overshadowing or facilitation of spatial pattern learning by visual cues

Two experiments were conducted to examine the effects of redundant and relevant visual cues on spatial pattern learning. Rats searched for hidden food items on the tops of poles that formed a square (Experiment 1) or a checkerboard (Experiment 2) pattern. The experimental groups were trained with visual cues that specified the locations of the baited poles. All groups were tested without visual cues so that any overshadowing or facilitation of spatial pattern learning by visual cues could be detected. Spatial choices were controlled by the spatial pattern and by the visual cues in both experiments. However, there was no evidence of overshadowing or facilitation of spatial pattern learning by visual cues in either experiment. The results are consistent with the idea that the representation of the spatial pattern that guides choices is not controlled by the same learning processes as those that produce associations between visual cues and food locations.     

Social working memory: Memory for another rat's spatial choices can increase or decrease choice tendencies

In two experiments using a radial-arm maze, pairs of rats made choices among eight maze locations, each containing a large quantity of one of two food types. The choices made by 1 rat affected the choices made by the other rat. Under most conditions, visits by 1 rat increased the tendency of the other rat to subsequently choose that maze location. However, the effect depended on the quality of the food available in a particular location. When it was possible for the rats to observe each other on the maze arms and a rat had experienced that a location contained the less preferred food type, a previous visit to that location by the foraging partner decreased the tendency to visit that location. These effects are attributed to working memory for the spatial choices of another rat, and they indicate that memory produced by a rat’s own visit to a maze location is integrated with memory for the behavior of another rat to determine spatial choice     

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.     

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.     

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.     

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.     

Two sources of proactive interference in spatial working memory: Multiple effects of repeated trials on radial maze performance by rats

Rats were trained in 8- and 12-arm radial mazes. Each trial began with a study phase (forced choices of 4 arms). The trial ended after a 2-h delay in a test phase consisting of free choices among 8 arms; choices of the 4 arms not yet visited were correct (rewarded). Proactive interference (PI) was induced by an interference phase that occurred on some days 2 or 3 h prior to the study phase. In the PI-repetition condition, the interference phase consisted of forced choices of the 8 arms that were later presented in the study and test phases; in the PI-nonrepetition condition, the interference consisted of forced choices of the 4 arms that were correct during the test phase. Test-phase performance was most accurate in the No PI (single-trial) condition and least accurate in the PI-repetition condition. A second experiment showed that repetitions per se were not responsible for the PI; when the interference phase consisted only of choices of the same 4 arms later presented in the study phase, no PI was observed. These findings suggest two sources of PI. One source, measured by the difference between No PI and PI-nonrepetition conditions, appears to be a difficulty in discriminating the temporal order of visits to arms in the interference and study phases. The other source of PI, measured by the difference between the nonrepetition and repetition conditions, remains to be identified; some possibilities are discussed.     

Time-place learning in the eight-arm radial maze

Rats (n=4) searched for food on an eight-arm radial maze. Daily 56-min sessions were divided into eight 7-min time zones, during each of which a different location provided food; locations were randomized across subjects before training. The rats obtained multiple pellets within each time zone by leaving and returning to the correct location. Evidence that the rats had knowledge about the temporal and spatial features of the task includes the following. The rats anticipated locations before they became active and anticipated the end of the currently active locations. The rats discriminated currently active locations from earlier and forthcoming active locations in the absence of food transition cues. After the rats had left the previously active location, they visited the next correct location more often than would be expected by chance in the absence of food transition cues. The rats used handling or opening doors as a cue to visit the first location and timed successive 7-min intervals to get to subsequent locations.     

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.     

Rats in a levered T-maze task show evidence of time–place discriminations in two different measures

It is difficult for rats to learn to go to an arm of a T-maze to receive food that is dependent on the time of day, unless the amount of food in each daily session is different. In the same task, rats show evidence of time–place discriminations if they are required to press levers in the arms of the T-maze, but learning is only evident when the first lever press is considered, and not the first arm visited. These data suggest that rats struggle to use time as a discriminative stimulus unless the rewards/events differ in some dimension, or unless the goal locations can be visited prior to making a response. If both of these conditions are met in the same task, it might be possible to compare time–place learning in two different measures that essentially indicate performance before and after entering the arms of the T-maze. In the present study, we investigated time–place learning in rats with a levered T-maze task in which the amounts of food varied depending on the time of day. The first arm choices and first lever presses both indicated that the rats had acquired time–place discriminations, and both of these measures became significantly different from chance during the same block. However, there were subtle differences between the two measures, which suggest that time–place discrimination is aided by visiting the goal locations.     

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.     

Schedule-induced wood-chewing in rats and its dependence on body weight

In Experiment 1, 12 rats were exposed to an FT 60 schedule of food reinforcement, followed either by extinction or by a massed-food control condition, in the presence of a wood block. In 9 rats, wood-chewing behavior increased systematically during the FT 60 condition and declined again during extinction or massed food, while the other 3 rats showed virtually no chewing behavior at any stage of the experiment. In Experiment 2, frequency and bout duration of wood-chewing under an FT 60 schedule of food reinforcement declined as body weight increased, in 7 rats. We conclude that wood-chewing qualifies as a schedule-induced behavior, and that it resembles schedule-induced drinking in its dependence on body weight. Unlike drinking, however, induced chewing occupied the middle region of the 60-sec interreinforcement interval, declined markedly within the session, and showed considerable within- and between-subject variability.     

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.     

Location, location, location: development of spatiotemporal sequence learning in infancy

We investigated infants' sensitivity to spatiotemporal structure. In Experiment 1, circles appeared in a statistically defined spatial pattern. At test 11-month-olds, but not 8-month-olds, looked longer at a novel spatial sequence. Experiment 2 presented different color/shape stimuli, but only the location sequence was violated during test; 8-month-olds preferred the novel spatial structure, but 5-month-olds did not. In Experiment 3, the locations but not color/shape pairings were constant at test; 5-month-olds showed a novelty preference. Experiment 4 examined "online learning": We recorded eye movements of 8-month-olds watching a spatiotemporal sequence. Saccade latencies to predictable locations decreased. We argue that temporal order statistics involving informative spatial relations become available to infants during the first year after birth, assisted by multiple cues.     

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.     

Spatial patterns and memory for locations

Rats obtained food from the tops of vertical poles in a 5 × 5 matrix of locations. On each trial, the baited locations formed one of the two possible exemplars of a checkerboard spatial pattern. During training, locations that had been visited earlier in the trial were indicated by a visual cue. Following training, performance with and without the visual cues was compared. Spatial choices were controlled by the checkerboard spatial pattern. The visual cues enhanced the ability of rats to avoid revisits of locations. However, the visual cues did not enhance control by the spatial pattern, as would be expected if the same spatial memories were involved in avoidance of revisits and coding the location of baited locations.     

Anticipatory contrast as a function of access time and spatial location

Intake of a 0.15% saccharin solution was suppressed when it was followed by a 32% sucrose solution in brief daily pairings. With equal access durations to the two solutions, intervals of intermediate duration (2 or 3 min) produced a larger contrast than more extreme intervals (1 or 10 min). There was no evidence of inhibition of delay with the 10-min interval (Experiments 1A and 1B). When access times were asymmetrical, longer access time to the first solution reduced contrast, whereas longer access time to the second solution enhanced contrast (Experiment 2). Contrast was greater when the two solutions were presented at consistent and separate spatial locations than when location was changed randomly or when both solutions were presented in sequence at the same location. However, a degree of contrast occurred in all conditions (Experiment 3). Experiment 4, conducted with the solutions in opposite arms of a T-maze, showed that anticipatory approach to the location correlated with the 32% sucrose solution developed prior to lick suppression on the saccharin solution. However, within daily sessions, there was a reliable increase in contrast without correlated changes in anticipatory-approach behavior. Access-time effects were attributed to altered reward values, whereas spatial-separation effects suggest that goal-directed responses contribute to, but do not cause, anticipatory contrast.     

Level of deprivation does not affect degree of discounting in pigeons

Two experiments tested the effects of food deprivation on discounting in pigeons. An adjusting-amount procedure was used to estimate the subjective value of food at delays ranging from 1 to 24 s. Experiment 1 compared pigeons’ discounting of delayed food reinforcers at 75 %–80 % and 90 %–95 % of free-feeding weight. Experiment 2 compared discounting under 1- and 23-h food deprivation. In both experiments at both deprivation levels, discounting was well described by the hyperboloid discounting function. No systematic effect of level of deprivation on degree of discounting was observed in either experiment. This finding is consistent with the view that pigeons’ choices are controlled by the relative, rather than the absolute, value of reinforcers.