Gerbils in space: performance on the 17-arm radial maze.

In Experiment 1 six hungry gerbils received six trials per day on a 17-arm radial maze. During each trial the subjects were allowed to choose freely among the arms, each of which contained a food pellet, until each arm had been visited once or until eight minutes had elapsed. An error was recorded when the subject entered a previously visited arm. The gerbils quickly learned not to re-enter previously visited arms and generally made errors on fewer than 15% of entries, performance comparable to that of the rat and superior to that of other species tested in the radial arm maze. The intertrial-interval duration did not affect accuracy of arm choices during acquisition but did influence asymptotic accuracy. Accuracy did not change systematically over the six trials. A high proportion of arm entries were to nearby arms. Errors occurred most often towards the end of a trial. Odor cues were not important. When the number of trials per day was reduced from six to one, accuracy deteriorated slightly. In Experiment 2 neither the transposition of extramaze cues nor the placement of the maze in a different room had large disruptive effects on accuracy. In Experiment 3 the addition of three explicit intramaze brightness cues aided accuracy, perhaps by permitting the subjects to decompose the large maze into three smaller mazes, although there was no direct evidence that this was the case. Implications of a number of these results for models of spatial maze performance were discussed.     

Spontaneous alternation behavior in hamsters

Male and female golden hamsters spontaneously alternated successive entries of T-maze arms (derived from a + − maze) rather than body turns during a series of eight consecutive trials except when not confined to the arm chosen on each trial. Only unconfined males failed to alternate on their first opportunity. Confinement to an entered arm for 30 seconds had no effect on females but increased alternation in males. Increasing the visual discriminability of the arms had no effect on alternation in either sex. It was concluded that spontaneous alternation behavior in hamsters depended upon whether or not they were confined to a chosen arm and their sex. It is possible that the phenomenon was escape-related.     

Transfer from discrimination under hunger to discrimination under thirst: The role of expectancy and habit

The experimental group (Group HA-TA) received food (F) and water (W) rewarded trials in an alternating sequence under hunger in Phase 1 and under thirst in Phase 2. Group HA-TA ran faster on F than on W trials in Phase 1, and faster on W than on F trials in Phase 2. Early in Phase 2 the difference between speeds on W and F trials was larger for Group HA-TA than for a group which received no runway training in Phase 1 (Group HO-TA), but later in Phase 2 this difference was larger for Group HO-TA than for Group HA-TA. Also in Phase 2 the difference between speeds on W and F trials was larger for Group HA-TA than for a group which received a random sequence of F and W trials under hunger in Phase 1, and smaller for Group HA-TA than for a group which received alternating F and W trials under thirst in both phases. To interpret these results it was assumed that for Group HA-TA the expectancies of reward formed in Phase 1 facilitated development of alternation performance in Phase 2, but that the S-R associative connections formed in Phase 1 inhibited ultimate development of alternation performance in Phase 2.     

Effects of scopolamine on repeated acquisition of radial-arm maze performance by rats.

Rats repeatedly acquired the performance of selecting only the four baited arms in an automated eight-arm radial maze, with the arms containing food pellets randomly assigned prior to each session. During each 14-trial (trial: obtain all four pellets) daily session, the number of errors (selecting nonbaited arms or repeating arm selections) showed a within-session decline, and choice accuracy for the first four arm selections showed a positive acceleration across trials for all rats. An index-of-curvature statistic, calculated for total errors, was used to quantify both the within- and between-session improvement of performance. Scopolamine (0.03 to 0.3 mg/kg, ip), but not methylscopolamine (0.3 mg/kg), reduced the accuracy of the first four selections of each trial and increased total within-session errors for all rats. Session times also were increased by scopolamine. An examination of within-session accuracy showed only slight signs of improvement at the higher dosages of scopolamine. The results indicate that behavior in transition states maintained by reinforcement contingencies in the radial maze is similar to that maintained by extended chained schedules, despite the fact that some of the stimuli controlling behavior in the maze are absent at the moment behavior is emitted.     

Differential Effects of Global Ischemia on Delayed Matching- and Non-Matching-to-Position Tasks in the Water Maze and Skinner Box

In order to assess effects of global ischemia in tasks of spatial learning and working memory, male Wistar rats were subjected to four vessel occlusion (4 VO) for periods of 5, 10, and 20 min and compared with sham-operated controls over four test phases, from 6 to 54 weeks after surgery. Rats were assessed on acquisition in the water maze, a task that is sensitive to ischemic impairments, before testing in Skinner box and water maze working memory tasks, which both require the short-term storage of information, but make different demands on spatial information processing. Phases 1 and 3 assessed spatial learning in a standard water maze procedure (12 and 10 training days, 2 trials/day with a 10-min intertrial interval: ITI). Phase 2 involved training and testing in delayed non-matching-to-position task in the Skinner box, with delays of 2–10 s between the information and choice stages. Phase 4 examined working memory in a water maze delayed matching-to-position task with 4 trials/day, an ITI of 30 s, and a novel platform position on each day. Ischemic rats showed duration-related impairments in water maze acquisition and working memory, but not in the less spatially demanding Skinner box task. Since water maze acquisition deficits were seen both before and after testing in the Skinner box the lack of effect cannot be attributed to time or to prior training. Ischemic deficits were more marked in Phase 3 than in Phase 1 of acquisition, suggesting that impairment may be progressive. Histological assessment showed that cell loss was largely confined to the hippocampal CA1 field and was linearly related to duration of occlusion. At the maximal level of loss (5.7 mm before the interaural line) the 20-min group showed 90% loss, the 10-min group 60% loss, and the 5-min group, which did not differ from controls, less than 10% loss. Only the 20-min group showed significant damage beyond the CA1 field, ranging from 30–40% loss in the CA3 field to 5% loss in one striatal area. No cortical damage was seen. The extent of CA1 cell loss correlated modestly with water maze acquisition (Phase 3) and working memory scores, but not with trials to criterion in the Skinner box task. There were significant correlations between different measures both within and between water maze tasks, but not Skinner box tasks, suggesting that the two types of procedure engaged different cognitive processes. The results indicate that the intrahippocampal damage induced by 4 VO impaired tasks which required processing of allocentric spatial information, but did not impair the storage of limited spatial information in working memory.     

Memory persistence of rats in a radial maze varies with training procedure

Different estimations of the time-course of spatial working memory have been reported. Some authors found working memory in the radial maze to be relatively long lasting, while others found more rapid exponential decay. In the present experiments it was attempted to account for the conflicting results by investigating the effects of different training procedures. Two types of training were examined under the same circumstances. A group of seven rats was given a series of delayed trials (5, 20, 60, and 120 min). Every delay was repeated at least four times and the delays were presented in an ascending order. The number of errors decreased at every delay except the last one (120 min), where error levels were constant (.50 errors/trial). The good performance was not based on use of intramaze cues or response chaining. In another group of seven rats the same delays were introduced in a quasi-random order and alternated with uninterrupted trials. The number of errors increased exponentially with the length of the delay. However, when this procedure was repeated, the number of errors decreased. These results suggest that training with delayed trials is a major factor to account for the differences in reports of memory persistence in the radial maze.     

Memory systems in the rat: effects of reward probability,context, and congruency between working and reference memory

The interaction of working and reference memory was studied in rats on an eight-arm radial maze. In two experiments, rats were trained to perform working memory and reference memory tasks. On working memory trials, they were allowed to enter four randomly chosen arms for reward in a study phase and then had to choose the unentered arms for reward in a test phase. On reference memory trials, they had to learn to visit the same four arms on the maze on every trial for reward. Retention was tested on working memory trials in which the interval between the study and test phase was 15 s, 15 min, or 30 min. At each retention interval, tests were performed in which the correct WM arms were either congruent or incongruent with the correct RM arms. Both experiments showed that congruency interacted with retention interval, yielding more forgetting at 30 min on incongruent trials than on congruent trials. The effect of reference memory strength on the congruency effect was examined in Experiment 1, and the effect of associating different contexts with working and reference memory on the congruency effect was studied in Experiment 2.     

The roles of beaconing and dead reckoning in human virtual navigation

Beaconing is a process in which the distance between a visual landmark and current position is reduced in order to return to a location. In contrast, dead reckoning is a process in which vestibular, kinesthetic and/or optic flow cues are utilized to update speed of movement, elapsed time of movement, and direction of movement to return to a location. In the present experiment, human participants engaged in a virtual triangle-completion task devoid of vestibular cues in which they navigated two legs of an outbound path before homing (returning to the origin). However, our task was modified such that a landmark array was placed at the origin, allowing both beaconing and dead reckoning to be utilized to home. During the second leg of the outbound path, we manipulated aspects of the landmark array to determine the extent to which beaconing and dead reckoning influenced homing. Specifically, during probe trials, the landmark array either remained the same, shifted 3 or 6 meters east/west, or north/south, or was removed. We analyzed homing performance to determine the influence of landmark shifts on search behavior. Results suggested that participants were able to home with equal accuracy in both the presence and the absence of the landmark array. Array shifts that put beaconing and dead reckoning in conflict resulted in an averaging of the two sources of information for the determination of heading but a mixture of strategies for path length.     

Effects of opiate antagonists on spatial memory in young and aged rats

The effects of post-training opiate antagonist administration on spatial memory were assessed in young and aged male Long Evans rats. In Experiment I rats were trained to visit each arm of an eight-arm radial maze once in a session to obtain a food reward placed at the end of each arm. During training aged rats required significantly more trials to achieve criterion performance when compared to young mature rats. However, administration of the opiate antagonist naloxone (2.0 mg/kg) immediately after each training trial did not significantly alter the rate of achieving accurate performance in either age group. In Experiment II young and aged rats that were previously trained to a comparable criterion on the radial maze were tested on the same maze apparatus in novel spatial environments. When animals were exposed to novel spatial information, the effects of post-trial opiate antagonists were examined using a within-subjects counter-balanced design. In Experiment IIa naloxone (2 mg/kg) enhanced the performance of both young and aged rats. In Experiment IIB naltrexone (1.0 mg/kg) was found to have a comparable effect of enhancing the performance of both age groups. In addition, in Experiment IIb a significant age-related deficit was found in rats tested in novel spatial environments. These results indicate that opiate antagonists are capable of improving memory for new spatial information in both young and aged rats on a task that is sensitive to behavioral deficits during normal aging.     

Establishing auditory stimulus control over an eight-member equivalence class via conditional discrimination procedures

Two eight-member equivalence classes of visual stimuli were established during three phases of a training program. In Phase 1, two training arrangements were compared. In one, 3 subjects were taught on different trials to select from a single pair of comparison stimuli (A1, A2) in response to eight sample stimuli that were trained in pairs (B1, B2; C1, C2; D1, D2; E1, E2). In the second arrangement, subjects were taught to select from four pairs of comparisons (B1, B2; C1, C2; D3, D2; E2, E2) in response to two samples (A1, A2). Training with the single pair of comparison stimuli resulted in the development of equivalence relations (B1C1, B2C2, D1B1, D2B2, B1E1, B2E2, C1D1, C2D2, C1E1, C2E2, D1E1, D2E2, and their reciprocals) between the sample stimuli without direct training of these relations. In the other training arrangement, these relations among the comparison stimuli developed in the performance of 1 subject only. In Phase 2, three new pairs of stimuli (F1, F2; G1, G2; H1, H2) were substituted for three of the original pairs (B1, B2; C1, C2; D1, D2) and the training arrangements for the groups were reversed. Following training, the performances that showed equivalence relations on the probes in the first phase also showed equivalence relations in the second phase. If such relations did not develop in the first phase, they did not do so in the second phase. In Phase 3, relations between stimuli across the two previous phases (e.g., B1F1, B2F2, B1G1, B2H2, C1F1, etc.) were investigated. The 4 subjects whose performances showed the development of these relations were taught to select one stimulus from each class (E1 and E2) in response to a verbal label (I1 and I2) and then were tested to see if the verbal label controlled responding to the remaining members of the class (e.g., I1A1, I2A2, I1B1, I2B2, etc.). For 3 subjects, this generalized control occurred; for the 4th, generalization occurred only after verbal training with a second pair of visual stimuli (F1 and F2). In retests several months later, these auditory-visual relations were found to be intact or, if not, were recovered without direct training.     

Changing behavior within session: cyclicity and perseverance produced by varying the minimum ratio of a variable-ratio schedule

Four pigeons repeatedly chose between a fixed-ratio (FR) 20 and a variable-ratio (VR) 40 schedule of reinforcement, in which the minimum ratio of the VR cycled within each session. The minimum ratio ascended and descended (ASCDESC), descended and ascended (DESCASC), or remained constant (unchanging). In Phase 1, 2 birds (Group 1) were exposed to ASCDESC series and 2 birds (Group 2) were exposed to the DESCASC series. Choice proportions changed with the cycling minimum ratio for Group 2 but not for Group 1. In Phase 2, Group 1 subjects were exposed to the DESCASC series and Group 2 subjects were exposed to the unchanging condition. Although Group 1's choice proportions appeared to be undifferentiated in Phase 2, Group 2's choice proportions continued to cycle for more than 100 sessions. Group 2 subjects were then moved to the ASCDESC series in the third phase, and choice proportions cycled with the minimum ratio as in the first phase. The descending portion of the series was the more powerful determinant of cyclicity. Response rates also changed with the minimum component ratio, a finding that goes against the claim of universality of a rise-and-fall within-session pattern of responding. That preference varied despite the constancy of the average ratio requirement suggests nonlinear averaging in quantitatively representing a variable schedule's value. The strong perseverance observed also lends support to a growing body of literature on history effects.     

Discrimination of what,when, and where: Implications for episodic-like memory in rats

We investigated the discrimination of what, when, and where in rats (n = 5) using an eight-arm radial maze. Rats received daily training consisting of forced-choice visits to four baited arms, one of which was randomly chosen each day to contain chocolate (Phase 1). In Phase 2, all eight arms were available. After a short (30 min) retention interval (RI), the four arms that were not available in Phase 1 provided food. After a long (4 h) RI, the four remaining arms plus the arm containing chocolate provided food (i.e., the chocolate arm replenished). The rats visited the chocolate location after the long RI more than after the short RI. Next, chocolate was paired with lithium chloride, and subsequent testing used the long RI. The rats visited the chocolate location less after the taste-aversion manipulation than in previous training, demonstrating knowledge of what, when, and where. Implications for episodic-like memory are discussed.     

Vicious-circle behavior in the Y maze

Laboratory rats were given escape training in both arms of a Y maze followed by extinction in one of four conditions: regular (no shock), punished (both arms containing a shock mid-segment), choice, or forced (one arm containing a shock mid-segment, one arm shock free). Subjects in the punished and forced conditions required more extinction trials than those in the regular extinction condition. The choice condition was intermediate and not significantly different from the others. Choice subjects avoided the shock by choosing the safe alley early and consistently during extinction, but some continued to run for many trials without further punishment. It was suggested that the number of punished trials early in extinction is related to persistence of running, and the results were discussed in relation to the Mowrer-Brown and the discrimination hypotheses.     

Visual dominance in a lateral plane motor learning task.

Visual dominance was investigated in a motor learning task with the criterion movement being in the lateral plane of the body. The criterion movement was a 10-in. abduction of the arm. All subjects received four presentation trials for the criterion movement in each of the following conditions: dominant rotated arm, dominant unrotated arm, nondominant rotated arm, and nondominant unrotated arm. Three independent groups of 10 college-age subjects differed according to sensory stimuli given during presentation trials. The Kinesthetic group was blindfolded for presentation trials. The Visual and Kinesthetic group was unblindfolded for presentation trials. The Alternating group was blindfolded on half of the presentation trials and unblindfolded on the other half. All subjects carried out five blindfolded reproduction trials for each of the four conditions. Absolute error for the length of the reproduced movements was measured and no significant difference between groups was found. This suggests that visual dominance is reduced in movements outside the frontal plane when focal vision is not used. Planned comparison testing indicated the Alternating group was significantly more accurate than the Visual and Kinesthetic group.     

Probability-reward preferences of rhesus monkeys

The relative preferences of rhesus monkeys for reward probability versus amount were investigated with procedures which contrasted general experience with specific instructions, and evaluated the relationship between probability-amount combinations and preference strength. Four stimulus objects, each signifying a different combination of reward frequency and amount (100% with one unit; 50% with two units; 33% with three units; or 25% with four units), were presented in pairs, one pair per daily session, with trial schedules providing the same amount of reward within each set of 12 trials. In Phase A, 4 monkeys (Group 1) were tested on the six choice-pairs with no preliminary training. In Phase B, Group 1 was joined by an additional 4 monkeys (Group 2), and each of the tasks was preceded by a demonstration of the relevant stimuli, one at a time, together with their associated probabilities and amounts. Group 1 animals developed preferences during Phase A for the more frequently rewarded objects, which persisted into Phase B, whereas Group 2 animals showed no preferences. This result indicates that preliminary instructions concerning the reward combinations associated with stimulus objects can prevent the development of a preference for greater probability over greater amount of reward, but cannot extinguish it once it has been formed or reestablished within the context of a particular task.     

Stimulus perseveration in a water maze following exposure to controllable and uncontrollable shock

When placed in a water-filled maze, mice display a pronounced preference for the illuminated over the nonilluminated arm of the maze. Exposure to inescapable shock increased the time spent in the illuminated arm of the maze, and decreased the frequency of entries into the nonilluminated arm. When animals that had received shock entered the nonilluminated arm they exhibited more activity per second than nonstressed animals. Controllability over the stressor enhanced the preference for the illuminated arm; however, the contribution of this variable was dependent on the number of shock trials mice received. Following 180 escapable or inescapable shock presentations the preference for the illuminated arm was enhanced. The propensity to approach the illuminated arm declined following a greater number (360) of escapable shock trials, while the preference for the illuminated arm did not decline in mice that received inescapable shock. Both escapable and inescapable shock were also found to produce a transient disruption of discrimination performance in a task where animals were required to emit a contraprepared response (swim to dark), whereas these treatments were without effect on performance of the highly prepared response of approaching the illuminated arm. It is provisionally suggested that enhancement of the perseveration represents an innate response to stressful stimuli, but as animals learn mastery over the response contingencies, the persistence in adopting such a response strategy wanes. Moreover, despite the differential effects of escapable and inescapable shock on the perseverative tendency, discrimination accuracy may not be differentially affected by these treatments in a task where acquisition progresses quickly and where explicit cues are associated with the correct and incorrect arms of the maze.     

Bilateral facilitation of motor control in chronic hemiplegia

The present study addressed the efficacy of concurrently moving both arms, with and without a load added to the uninvolved arm, in facilitating the quality of movement of the involved side in individuals with moderate, chronic hemiplegia. Six hemiplegic cerebrovascular accident (CVA) subjects with left-hemisphere lesions participated in the study. The four males and two females ranged from 46 to 77 years of age and 30–96 months post-CVA. All subjects scored at least 70% on the Fugl-Meyer test of motor function. The task was to perform discrete unilateral and bilateral elbow extensions in the horizontal plane. The movements were 45° in amplitude and were to terminate in a 10° target zone that was indicated by an illustration of a coffee mug. The instructions were to move toward the mug as smoothly as possible in a movement time (MT) determined to be 20% longer than their minimal MT for that distance. The primary dependent variable was the percentage of continuous vs. discontinuous trajectories observed in each condition, based on whether or not a transient hesitation or reversal was observed. Phase of peak velocity was also quantified as a general indication of the symmetry of the velocity profile. Three of the six subjects exhibited a greater percentage of continuous movements of the involved arm in the nonloaded bilateral condition than the unimanual condition. Five subjects benefited when the uninvolved arm was inertially loaded in the bilateral condition when compared with unimanual performance. Only the oldest subject failed to exhibit facilitation. Peak velocity phase tended to normalize toward symmetry in the bilateral conditions. These findings are consistent with prior evidence that the control of the involved arm improves during bimanual performance for some hemiplegic subjects. It further suggests loading the uninvolved arm may benefit some subjects with respect to unimanual performance, with age perhaps playing a role in determining efficacy.     

Chunking by Proximal Arm Cues Facilitates Rats′ Performance in the Radial Maze

Three experiments showed that training rats to chunk a 16-arm radial maze into striped and wire mesh (gridded) arms facilitates their spatial working memory for arm locations. Rats were trained to visit eight unblocked baited arms of a 16-arm radial maze (half-maze run) and then were exposed to the complete maze to sample the remaining eight baited arms (whole-maze run). The initially exposed eight arms were either all striped or all gridded on alternating trials for half the rats (Arm Cue Relevant, ACR group). The remaining rats (Arm Cue Irrelevant, ACI group) received a mixture of striped and gridded arms on their half-maze runs. Following this phase of segmented trials, all rats were exposed only to all 16 arms over a series of trials in the first two experiments. In the third experiment, the initial eight arms were either all striped or all gridded on alternating trials for all rats. During some second, whole-maze runs, however, all arms contained the same proximal cue. ACR rats made fewer reentries than ACI rats in all phases of all experiments. This difference was maintained over increasing delays between half- and whole-maze runs in the first experiment, changes in arm cue and blocking configurations in the second experiment, and removal of differential arm cues in the third experiment.     

Learning about water by hungry rats

In Experiments 1 and 2 hungry rats were trained in a straight alley with consistent food reward (FF), food and nonreward trials intermixed (FN), or food and water trials intermixed (FW). In Experiment 1 rats were tested with nonrewarded trials (extinction) and Groups FN and FW did not differ, both running faster then Group FF. In Experiment 2 rats were tested with consistent water reward, and Group FW ran faster then Group FN, which was superior to Group FF. In Experiment 3, one group of hungry rats was trained on a single alternating schedule of food and water in Phase 1 and was shifted to a single alternating schedule of food and nonreward in Phase 2 (Group FW); the second group (Group FN) received a single alternating schedule of food and nonreward in both phases. When Group FW was shifted to nonreward from water, performance to nonreward was temporarily disrupted. These results indicate that, contrary to previous conclusions, hungry rats can learn about water when drive is strong and food is received in the situation.     

Recognition of a 3D snake model and its 2D photographic image by captive black tufted-ear marmosets (Callithrix penicillata)

Two-dimensional (2D) displays of real three-dimensional (3D) objects are frequently used experimental tools in animal studies. Whether marmoset monkeys, with their highly diverse and complex anti-predation strategies, readily recognized 2D representations of potential threats has yet to be determined, as seen in other primates. Thus, the behavioral responses of adult captive black tufted-ear marmosets (Callithrix penicillata) toward an unfamiliar motionless snake-model and its photograph were assessed. Pictorially naïve subjects were randomly divided into two groups (n = 12/each) and submitted to two trials. Group 1 was initially exposed to the 3D object and after 1 week to its photograph. Group 2 was first presented the picture and only tested with the real object 1 week later. All 15-min trials were divided into three consecutive 5-min intervals: pre-exposure, exposure and post-exposure. In the presence of the 3D snake object, regardless of its presentation order, the frequency of direct gazes, head-cocks, tsik-tsik alarm/mobbing calls and genital displays increased significantly. The photograph induced a similar response, although only when the object had been previously presented, as significantly higher levels of these behaviors were seen in Group 1 than Group 2. Proximity to the stimulus, aerial scan, terrestrial glance, displacement activities and locomotion were not consistently influenced by the stimuli’s presence and/or order of presentation. Therefore, marmosets recognized and responded appropriately to biologically and emotionally relevant 3D and 2D stimuli. Since the aversive/fearful reactions toward the photograph were only seen after the snake object had been presented, the former seems to be essentially a learned response.