Training delays reduce the choose-short effect with keylight duration samples in pigeons

Pigeons were trained to matching 2- and 8-s keylight samples. The delay on training trials was either 0s (group 0sF), 2s (group 2sF), or varied between 1 and 3s (M=2s, group 2sV). Testing at delays of 10 and 20s revealed a choose-short tendency in all three groups. The magnitude of this tendency was largest in group 0sF and was highly similar in magnitude in groups 2sF and 2sV. In Experiment 2, the training delay remained at 0s in group 0sF and was increased to 5s in group 2sF (now group 5sF). Group 2sV (now group 5sV) received variable training delays ranging from 2 to 8s (M=5s). Testing at a 20-s delay and at a delay that exceeded the training delay by 15s for each group revealed a robust choose-short effect only in group 0sF. Groups 5sF and 5sV both demonstrated a weak and statistically nonsignificant choose-short tendency. When different durations of keylight are employed as samples, training with a nonzero delay, whether fixed or variable, reduces the magnitude of the choose-short effect, with longer training delays producing a greater reduction in the choose-short effect.     

Effect of training delays and start and stop markers on the choose-short effect in pigeons

Pigeons were trained in a matching-to-duration task using short and long filled intervals. Group 2/8 was trained with 2- and 8-s intervals, Group 4/10 with 4- and 10-s intervals, and group marker with 2- and 8-s intervals presented between 1-s start and stop markers. Extended-delay testing showed no significant choose-short effect (CSE) in any group. It was hypothesized that the lack of a CSE may have resulted from use of a variable delay (range 1-3s) during training. The same subjects were employed in Experiment 2 and were trained with a new set of comparisons and one of the alternate types of samples employed in Experiment 1. All training trials involved a 0-s delay. Extended-delay testing revealed a significant CSE in Groups 2/8 and 4/10 but only a weak, and statistically nonsignificant, CSE in group marker. It was concluded that use of a variable delay during training reduced the CSE. The notion that subjective shortening underlies the CSE provided an adequate account of these findings.     

Choose-long errors at delays shorter than the training delay persist when enhanced between-trial summation of duration memories are precluded in pigeons

Pigeons were trained to match 2- and 10-s durations of houselight to red and green comparisons. Following acquisition with a 0-s baseline delay, they were tested with delays of 0, 10 and 20 s. There was a strong tendency to choose the short-associated comparison stimulus at both the 10- and 20-s delays (i.e., a choose-short effect) and no bias at the 0-s delay. This test was repeated after baseline training with a constant 10-s delay. As in the first delay test, a choose-short effect was obtained at the 20-s delay. In contrast to the first test, no bias was obtained at the 10-s delay and a strong tendency to choose the long-associated comparison (i.e., a choose-long effect) was obtained at the 0-s delay. Importantly, the choose-long effect was obtained under conditions which insured that the temporal spacing between a 0-s delay trial and the preceding trial was equal to that in training. These results are inconsistent with the temporal summation account of the choose-long effect and are most readily interpreted within a perspective emphasizing the subjective shortening of temporal memories over time.     

Effects of delays on human performance on a temporal discrimination procedure: evidence of a choose-short effect

Studies of temporal discrimination in non-human subjects have reliably shown a choose-short effect: higher matching accuracy on short-duration-sample trials than on long-duration-sample trials. This effect occurs as a function of increasing the delay between the onset of sample and comparison stimuli in a delayed matching-to-sample procedure. The present experiment investigated whether the choose-short effect could be demonstrated in human subjects under conditions which paralleled those used with non-human subjects. Subjects responded under a discrete-trial procedure in which they were required to push one of two buttons depending on the duration of a sample stimulus (a blue square on a computer monitor). Delays (0, 8, 16, and 32s) separated sample and comparison stimuli (two white boxes) and were tested both within and across several sessions. Intermediate durations (probe stimuli between 2 and 4s) were also presented. The addition of a delay between the sample and comparison stimuli produced a bias to judge intervals as short when the 8 and 32-s delays were tested across sessions and when the 0, 16, and 32-s delays were tested within the same session. Thus, the choose-short effect was produced in human subjects using the interval bisection procedure regardless of delay length.     

The effects of concurrent task and gap events on peak time in the peak procedure

The effect of a concurrent task on timing performance of pigeons was investigated with the peak interval procedure. Birds were trained to peck a side key on a discrete-trial schedule that included reinforced fixed-interval (FI) 30-s trials and nonreinforced extended probe trials. Then, in separate sessions, birds were trained to peck a 6-s center key for food. In a subsequent test phase, the FI procedure was in effect along with dual-task probe test trials. On those test trials, the 6-s center key (task cue) was presented at 3, 9, or 15s after probe trial onset. During another test phase, a 6-s gap (the FI keylight was extinguished) was presented at 3, 9, or 15s after probe trial onset. Peak time increased with center key time of onset, and was greater under task than gap conditions. Moreover, peak time under task conditions exceeded values predicted by stop and reset clock mechanisms. These results are at variance with current attentional accounts of timing behavior in dual-task conditions, and suggest a role of nontemporal factors in the control of timing behavior.     

The choose-short effect in rat memory for event duration: the subjective-shortening model

Two predictions derived from the subjective-shortening model were tested in rats. The predictions concerned the temporary occurrence of the choose-short effect with extended training at a given retention interval (RI) and the occurrence of a temporary choose-long effect, when RIs shorter than those used during training were applied. In a first experiment, using a stepwise delay procedure with training 0-s RI sessions interpolated between each series of increasing RIs, results showed: (1) a choose-short effect during the stepwise increase in the delay procedure, (2) a temporary occurrence of the choose-short effect during testing at a given RI and (3) a choose-long effect in half of the animals, when a RI shorter than that used previously was applied. These contrasting results suggest that the disappearance of the choose-short effect could be, as proposed by the model, either the consequence of the foreshortening of the reference memory (for rats choosing-long) or the consequence of an adaptation of the working memory (for rats which did not choose long). Results were discussed in relation with the procedure which could have contributed, by the interposition of 0-s RI sessions, to maintain a stable reference memory. In order to test this interpretation, a second experiment, using the classical stepwise delay procedure without training sessions interpolated, was carried out. In these conditions, rats did never present a significant choose-long effect when the RI was shortened. These results suggest that rats maintained a stable reference memory and could improve their performances during retention testing sessions either by an adaptation of their working memory or by the adoption of an alternative strategy which consisted in learning to maintain an orientation towards the location of the correct lever.     

The effects of cue competition on timing in pigeons

We studied the effects of cue competition on timing in both overshadowing and blocking operant procedures with pigeons. A white center key delivered reward when pecked 30 s after a red or green sidekey was presented and 10 s after presentation of the alternate color on the other sidekey. In Experiment 1, key presentations were concurrent during training trials for overshadow-condition pigeons, while side key presentations were separated across training trials for control birds. In Experiment 2, half of the birds (Blocking group) were given pre-exposure trials to either the 10-s or 30-s sidekey condition. Both blocking-condition and control birds were then given trials of concurrent side key presentations. Peak time curves were compared between experimental and control conditions. The results showed blocking of timing accuracy of a long (30-s) stimulus by a short (10-s) stimulus, but no evidence for overshadowing of timing accuracy.     

Effects of retention interval on performance in a numerical reproduction task

The retention interval (RI) between the sample and production phase in a numerical reproduction task was varied to determine whether a "produce-small" effect would be obtained with increased delays. Four pigeons were trained with a retention interval of 2s, and then tested with intervals of 0.5s and 8s. Results showed a number-dependent, "produce-large" effect-response number increased when RI was increased-analyses of average response number and accuracy suggested RI affected responding most on the 2-flash trials with an 8-s RI. Additionally, discrimination between trial types decreased as RI increased. Existing explanations for the "choose-short/small" effect appear unable to account for these results; however the "produce-large" effect may be attributed to a disruption in stimulus control over responding.     

Effect of US identity on elimination and recovery of autoshaped responding with explicitly unpaired and degraded contingency extinction procedures

The depressive effects of noncontingent and explicitly unpaired food unconditioned stimuli (USs) and the recovery from those effects on autoshaped responding were examined in a series of experiments with pigeons. In Experiments 1 and 2, responding to a keylight conditioned stimulus (CS) previously paired with food was depressed equally by explicitly unpaired presentations of either that same food or a different food. Furthermore, responding recovered equally following removal of the explicitly unpaired foods. In contrast, Experiments 3 and 4 showed that noncontingent presentations of a food US depressed responding more to a keylight CS paired with that same food than to a keylight CS paired with a different food. Moreover, removal of the noncontingent foods led to complete recovery but more rapid extinction of responding to the same keylight relative to the different keylight. The implications of these results for understanding the mechanisms underlying depression and recovery of responding following degraded contingency and explicitly unpaired extinction procedures are discussed.     

Context effect in a temporal bisection task with the choice keys available during the sample

In a symbolic matching to sample task, six pigeons learned to associate different sample durations with different comparison stimuli. On “Short” trials, choice of Red and Green keylight comparisons were reinforced following 3-s and 9-s samples, respectively; on “Long” trials, Blue and Yellow keylight comparisons were reinforced following 9-s and 27-s samples, respectively. In contrast with previous studies, the comparison keys were available during the samples. After the temporal discriminations were learned, new pairs of comparison keys were presented and the preference for each was assessed during 27-s samples. One pair in particular, Green and Blue, was critical because it tested the predictions of two timing models, Scalar Expectancy Theory (SET) and the Learning-to-Time (LeT) model. The results showed that preference for Green increased during the sample, a result consistent with LeT but not with SET. Other test results, however, were predicted by neither model.     

Interstimulus interval as a discriminative stimulus: Evidence of the generality of a novel asymmetry in temporal discrimination learning

Three appetitive conditioning experiments with rats examined temporal discrimination learning within Pavlovian conditioning trials. In all experiments, the duration of a feature white noise stimulus signaled whether or not a subsequent 10-s target tone would be reinforced. In Experiment 1, the feature durations were 4 and 1 min. For one group of rats (Group 4+/1−), 4 min of noise signaled that the tone would be reinforced and 1 min of noise signaled that the tone would not be reinforced. A second group (Group 1+/4−) was trained with the reverse contingency. The results showed a clear asymmetry in temporal discrimination learning: rats trained with 4+/1− (Long+/Short−) learned the discrimination readily (responding more in the tone on reinforced than on nonreinforced trials), whereas rats trained with 1+/4− (Short+/Long) did not. In Experiment 2, the feature durations were shortened to 60 and 15 s. Due to strong excitatory conditioning of the 15-s feature, the reverse asymmetry was observed, with the Short+/Long− discrimination learned more readily than the Long+/Short− discrimination. However, Experiment 3 demonstrated that the original Long+/Short− advantage could be recovered while using 60− and 15-s feature durations if the excitatory conditioning of the feature was reduced by including nonreinforced feature trials. The results support previous research involving the timing of intertrial intervals and are consistent with the temporal elements hypothesis which holds that the passage of time is encoded as a series of hypothetical stimulus elements.     

Serial conditional discrimination and temporal bisection in rats selectively lesioned in the dentate gyrus

In positive serial conditional discrimination, animals respond during a target stimulus when it is preceded by a feature stimulus, but they do not respond when the same target stimulus is presented alone. Moreover, the feature and target stimuli are separated from each other by an empty interval. The present work aimed to investigate if two durations (4 or 16 s) of the same feature stimulus (light) could modulate the operant responses of rats to different levers (A and B) during a 5-s target stimulus (tone). In the present study, lever A was associated with the 4-s light, and lever B was associated with the 16-s light. A 5-s empty interval was included between the light and the tone. In the same training procedure, the rats were also presented with the 5-s tone without the preceding light stimuli. In these trials, the responses were not reinforced. We evaluated the hippocampal involvement of these behavioral processes by selectively lesioning the dentate gyrus with colchicine. Once trained, the rats were submitted to a test using probe trials without reinforcement. They were presented with intermediate durations of the feature stimulus (light) to obtain a temporal bisection curve recorded during the exposure to the target stimuli. The rats from both groups learned to respond with high rates during tones preceded by light and with low rates during tones presented alone, which indicated acquisition of the serial conditional discrimination. The rats were able to discriminate between the 4- and 16-s lights by correctly choosing lever A or B. In the test, the temporal bisection curves from both experimental groups showed a bisection point at the arithmetic mean between 4 and 16 s. Such processes were not impaired by the dentate gyrus lesion. Thus, our results showed that different durations of a feature stimulus could result in conditional properties. However, this processing did not appear to depend on the dentate gyrus alone.     

Temporal discrimination learning by pigeons

Memory for time by animals appears to undergo a systematic shortening. This so-called choose-short effect can be seen in a conditional temporal discrimination when a delay is inserted between the sample and comparison stimuli. We have proposed that this temporal shortening may result from a procedural artifact in which the delay appears similar to the intertrial interval and thus, produces an inadvertent ambiguity or 'instructional failure'. When this ambiguity is avoided by distinguishing the intertrial interval from the delay, as well as the samples from the delay, the temporal shortening effect and other asymmetries often disappear. By avoiding artifacts that can lead to a misinterpretation of results, we may understand better how animals represent time. An alternative procedure for studying temporal discriminations is with the psychophysical bisection procedure in which following conditional discrimination training, intermediate durations are presented and the point of subjective equality is determined. Research using the bisection procedure has shown that pigeons represent temporal durations not only as their absolute value but also relative to durations from which they must be discriminated. Using this procedure, we have also found that time passes subjectively slower when animals are required to respond to the to-be-timed stimulus.     

Rats' memory for event duration in delayed matching-to-sample with nonspatial comparison response alternatives

Previous research has suggested that using stationary and moving levers as nonspatial response alternatives can significantly enhance the speed of acquiring a temporal discrimination in rats. In Experiment 1, rats were trained to discriminate 2 and 8s of magazine light illumination by responding to either a stationary lever or a moving lever with a cue light illuminated above it. Rats learned to discriminate event durations at a high level of accuracy after 25 sessions of training. During subsequent delay tests, rats exhibited a strong choose-long bias, indicating that they were timing from the onset of the magazine light until the entry of levers into the chamber. This occurred regardless of whether intertrial intervals and delay intervals were dark or illuminated. On test trials in which the sample was omitted, rats responded as if the short sample had been presented. In Experiment 2, the rats received extensive training with dark and illuminated variable delay intervals (1-4 s). However, they continued to exhibit a tendency to time from the onset of the magazine light until entry of the levers into the chamber. Although the use of stationary/uncued and moving/cued levers as response alternatives enhanced the speed of acquisition of the event duration discrimination in rats, additional procedural modifications will be necessary to prevent rats from timing during the delay interval.     

Representation of time intervals in a double bisection task: Relative or absolute?

We examined if the representation of time intervals in a temporal discrimination task is based not only on their absolute but also on their relative durations. Six pigeons learned two temporal discriminations. In the first, red and green choices were correct following 2-s and 8-s samples, respectively. In the second, vertical and horizontal bar choices were correct following 4-s and 16-s samples, respectively. In a previous study [Zentall, T.R., Weaver, J.E., Clement, T.S., 2004. Pigeons group time intervals according to their relative duration. Psychon. Bull. Rev. 11, 113-117.], tests with 4-s samples and red/green comparisons revealed a bias for red, whereas tests with 8-s samples with vertical/horizontal comparisons revealed a bias for horizontal. These results were interpreted in terms of relative encoding of sample durations. We attempted to replicate this finding but instead of testing with only 4-s or 8-s samples, we tested with several other sample durations to obtain a psychometric function. Results were inconsistent with the relative encoding hypothesis.     

The effects of changes in consequences on hens' performance in delayed-matching-to-sample tasks

A delayed-matching-to-sample (DMTS) task was used to investigate remembering with domestic hens. In Conditions 1 and 3 of Experiment 1, six hens responded under a mixed-delay procedure with delays of 0.25, 2, and 8 s. In Condition 2, the reinforcer for correct responding was delayed for 6 s after each correct matching response on 2-s delay trials. In Condition 1, discrimination performance decreased monotonically over the three delays. With the delay to the reinforcer, the decreases were non-monotonic as a result of the considerable drop in the accuracy of discrimination on the 2-s delay trials. Performance at the 2-s delay did not recover completely in Condition 3. In Conditions 1 and 3 of Experiment 2, five hens responded under a mixed-delay procedure with delays of 0, 4, and 16 s. In Condition 2 no reinforcers were provided for correct responding on 0- and 16-s delay trials. When reinforcers were available on all trials discrimination performance decreased monotonically with delay. There were non-monotonic changes in discrimination with delay when there was extinction at two delays resulting mainly from a large drop in discrimination performance at 0 s. In addition, response latencies increased markedly at the two delays associated with extinction. Performance recovered completely in Condition 3. The data support the ideas that remembering involves a temporal discrimination that the effects of delaying reinforcement and removing reinforcement may differ, and that the measurement of response latencies may be a useful tool in DMTS procedures.     

Effects of unsignaled delays of reinforcement on fixed-interval schedule performance

Key pecking of pigeons was maintained by a fixed-interval (FI) 61-s schedule. The effects of resetting and nonresetting unsignaled delays of reinforcement then were examined. The resetting delay was programmed as a differential-reinforcement-of-other-behavior schedule, and the nonresetting delay as a fixed-time schedule. Three delay durations (0.5, 1 and 10 s) were examined. Overall response rates were decreased by one and 10-s delays and increased by 0.5-s delays. Response patterns changed from positively accelerated to more linear when resetting or nonresetting 10-s delays were imposed, but remained predominantly positively accelerated when resetting and nonresetting 0.5- and 1-s delays were in effect. In general, temporal control, as measured by quarter-life values, changed less than overall response rates when delays of reinforcement were in effect. The response patterns controlled by FI schedules are more resilient to the nominally disruptive effects of delays of reinforcement than are corresponding overall response rates.     

Retention deficit for avoidance training in hypophysectomized rats: Time-dependent enhancement of retention performance with post-training ACTH injections

These experiments examined the effects of hypophysectomy on retention of avoidance training. In addition, the experiments examined the effects, on retention, of post-training ACTH injections administered to hypophysectomized rats. Rats were trained in a visual discriminated avoidance Y maze. Each rat received six training trials followed by six retraining trials the next day. Retention was measured by the number of correct choices during the retraining trials. When trained with a low-footshock intensity (0.8 mA), hypophysectomized rats showed retention performance which was significantly poorer than that of intact animals. There was no significant difference in performance when the animals were trained with a higher footshock intensity (1.4 mA), in part because of poorer retention performance of intact animals under these training conditions. Under both footshock conditions, a single post-training injection of ACTH enhanced later retention performance of hypophysectomized rats. This effect on memory was timedependent; injections delayed 2 or 6 hr after training did not significantly enhance retention. These findings are consistent with the view that hormonal responses to training may modulate later retention of the training experience.     

Nicotine and the behavioral mechanisms of intertemporal choice

Nicotine has been found to produce dose-dependent increases in impulsive choice (preference for smaller, sooner reinforcers relative to larger, later reinforcers) in rats. Such increases could be produced by either of two behavioral mechanisms: (1) an increase in delay discounting (i.e., exacerbating the impact of differences in reinforcer delays) which would increase the value of a sooner reinforcer relative to a later one, or (2) a decrease in magnitude sensitivity (i.e., diminishing the impact of differences in reinforcer magnitudes) which would increase the value of a smaller reinforcer relative to a larger one. To isolate which of these two behavioral mechanisms was likely responsible for nicotine's effect on impulsive choice, we manipulated reinforcer delay and magnitude using a concurrent, variable interval (VI 30 s, VI 30 s) schedule of reinforcement with 2 groups of Long-Evans rats (n = 6 per group). For one group, choices were made between a 1-s delay and a 9-s delay to 2 food pellets. For a second group, choices were made between 1 pellet and 3 pellets. Nicotine (vehicle, 0.03, 0.1, 0.3, 0.56 and 0.74 mg/kg) produced dose-dependent decreases in preference for large versus small magnitude reinforcers and had no consistent effect on preference for short versus long delays. This suggests that nicotine decreases sensitivity to reinforcer magnitude.     

Pigeons' memory for sequences of light flashes: reliance on temporal properties and evidence for delay interval/gap confusion

In Experiment 1, pigeons were trained to discriminate between sequences of two and four light flashes (illumination of the feeder). Retention functions obtained with dark delays exhibited a choose-many bias at a 1-s delay and a choose-few bias at delays of 4 and 8s. Retention functions obtained with illuminated delays only displayed a slight choose-few bias. In Experiment 2, additional birds were trained with the same sample sequences. However, the intertrial interval was illuminated by the houselight for Group Light, and it was dark for Group Dark. The acquisition data suggested that multiple temporal features of the light flash sequences controlled choice responding. For both groups, the retention functions were similar to those obtained in Experiment 1. In Experiment 3, baseline training with a 1-s dark delay eliminated the choose-many bias, but a significant choose-few bias at extended dark delays was still observed. Pigeons discriminate light flash sequences by relying on temporal properties of the sequence rather than using an event switch to count flashes. The biased-forgetting effects obtained in these studies appear to be primarily due to confusion between the delay interval and the gap between light flashes.