Comparison of timing and classical conditioning

Four experiments with rats investigated if the timing of a stimulus (sound) correlated with the strength of a conditioned response (CR) to the stimulus. The timing (effective duration) of the stimulus was measured using the peak procedure, similar to a discrete-trials fixed-interval procedure. The rats were trained so that their response rate reached a maximum about 40 s or 60 s after the onset of a light; the time of the maximum measured from the start of the light (peak time) was the measure of timing. On some trials, the light was preceded by a short (5 s) or long (20 s or 30 s) interval of sound. We assumed that the difference in peak time after long and short sounds reflected the timing of the sound--if the sound was timed, the longer sound would produce a lower peak time; if the sound was not timed, the two durations of sound would produce the same peak time. The CR was lever-pressing during the sound. The sound was treated in various ways: presented alone (Experiments 1, 3, and 4), followed by food (Experiments 1, 3, and 4), preceded by food (Experiment 3), and followed by food after 20 s (Experiment 4). Treatments that produced no timing of sound produced no CR, and treatments that increased (or diseased) timing also increased (or decreased) the CR. The results suggest that there is overlap between the mechanisms that produce time discrimination and the mechanisms that produce classical conditioning.     

Peak Procedure Perform ance in Young Adult and Aged Rats: Acquisition and Adaptation to a Changing Temporal Criterion

Twenty-four-month-old and 4-month-old rats were trained on a peak-interval procedure, where the time of reinforcement was varied twice between 20 and 40 sec. Peak times from the old rats were consistently longer than the reinforcement time, whereas those from younger animals tracked the 20- and 40-sec durations more closely. Different measures of performance suggested that the old rats were either (1) systematically misremembering the time of reinforcement or (2) using an internal clock with a substantially greater latency to start and stop timing than the younger animals. Old rats also adjusted more slowly to the first transition from 20 to 40 sec than did the younger ones, but not to later transitions. Correlations between measures derived from within-trial patterns of responding conformed in general to detailed predictions derived from scalar expectancy theory. However, some correlation values more closely resembled those derived from a study of peak-interval performance in humans and a theoretical model developed by Cheng and Westwood (1993), than those obtained in previous work with animals, for reasons that are at present unclear.     

Timing light and tone signals in pigeons

Pigeons' ability to time light and tone stimuli was examined in four experiments. In Experiment 1, two groups of pigeons were trained to discriminate between 2- and 8-s durations of lights or tones and then were transferred to reversal or nonreversal discriminations in the alternate modality. Pigeons learned the light discrimination faster than the tone discrimination and showed immediate positive intermodal transfer from tone to light but not from light to tone. In Experiments 2-4, the peak procedure was used to study birds' timing of 15- and 30-s fixed-interval light and tone signals. Peak times on empty trials under baseline conditions closely approximated the length of fixed-interval signals. When pigeons were tested with time-outs and intermodal switches introduced midway through an empty trial, they tended to reset the timing mechanism and begin timing again from 0 s. With both estimation and production procedures, pigeons were less accurate when timing the tone stimuli than when timing the light stimuli. A comparison of these data with data from timing experiments with rats suggests several possible differences in timing processes between pigeons and rats.     

The effects of aging on time reproduction in delayed free-recall

The experiments presented here demonstrate that normal aging amplifies differences in time production occurring in delayed free-recall testing. Experiment 1 compared the time production ability of two healthy aged groups as well as college-aged participants. During the test session, which followed a 24-h delay and omitted all feedback and examples of the two target intervals, the two groups of aged participants' over-produced a 6s interval. This effect is similar in form to errors shown by young participants, but twice the magnitude. Productions of a 17 s interval were generally accurate overall. However, further analysis indicated that the majority of aged participants over-produced the 17 s interval while a minority greatly under-produced the 17 s interval. Furthermore, aged participants showed violations of the scalar property of timing variability in the training session, while in the test session, only those who under-produced the 17 s interval showed this tendency. In contrast, training session performance was good for all participants. Experiments 2 and 3 investigated the ability of the participants in Experiment 1 to reproduce the length of a line from memory, under conditions analogous to those of the time production experiments. These experiments provided tests of the specificity of the errors observed in Experiment 1. Performance in the older participants was accurate, if more variable, compared to the young participants, in contrast to the time production results, indicating that production inaccuracy in free-recall is specific to interval timing in the current context.     

The power law and Weber's law in fixed-interval postreinforcement pausing: A scalar timing model

Some quantitative properties of the postreinforcement pause under fixed-interval schedules were simulated by a computer model embodying two processes, either of which could initiate responding in an interval. The first was a scalar timing system similar to that hypothesized to underlie behaviour on other tasks. The second was a process that initiated responding without regard to elapsed time in the interval. The model produced simulated pauses with a mean that varied as a power function of the interval value, and a standard deviation that appeared to grow as a linear function of the mean. Both these features were found in real data. The model also predicted several other features of pausing and responding under fixed-interval schedules and was also consistent with the results produced under some temporal differentiation contingencies. The model thus illustrated that behaviour that conformed to the power law could nevertheless be reconciled with scalar timing theory, if an additional non-timing process could also initiate responding.     

Representation of time

Memory representation for time was studied in two settings. First, an analysis of timing in a laboratory analog of a foraging situation revealed that departure times from a patchy resource followed a Weber Law-like property implied by scalar timing. A trial-by-trial analysis was then pursued in a similar but more structured experimental paradigm, the Peak procedure. Study of covariance structures in the data implicated scalar variance in the memory for time as well as in the decision process, but the correlation pattern ruled out multiple access to memory within a trial.     

Multiple-interval timing in rats: Performance on two-valued mixed fixed-interval schedules

Three experiments studied timing in rats on 2-valued mixed-fixed-interval schedules, with equally probable components, Fixed-Interval S and Fixed-Interval L (FI S and FI L, respectively). When the L:S ratio was greater than 4, 2 distinct response peaks appeared close to FI S and FI L, and data could be well fitted by the sum of 2 Gaussian curves. When the L:S ratio was less than 4, only 1 response peak was usually visible, but nonlinear regression often identified separate sources of behavioral control, by FI S and FI L, although control by FI L dominated. Data were used to test ideas derived from scalar expectancy theory, the behavioral theory of timing, and learning to time.     

Reward value effects on timing in the peak procedure

Three experiments examined the effect of motivational variables on timing in the peak procedure. In Experiment 1, rats received a 60-s peak procedure that was coupled with long-term, between-phase changes in reinforcer magnitude. Increases in reinforcer magnitude produced a leftward shift in the peak that persisted for 20 sessions of training. In a final phase, the rats received lithium chloride-induced aversion prior to testing and a rightward shift in the peak was observed. Experiment 2 confirmed the rightward shift in the peak under lithium chloride devaluation and induced a comparable shift with satiety devaluation. The degree of rightward shift was neither additive nor multiplicative, suggesting that two processes may have contributed. Experiment 3 examined the effect of extinction on peak responding, revealing a decrease in response rate, but no evidence of any change in the timing of responding. The implications of the results for contemporary timing theories are discussed.     

Simultaneous temporal processing

Seven experiments assessed the ability of rats to process temporal information from two internal clocks simultaneously and independently. In the first six experiments a light stimulus signalled an overall interval between the beginning of a trial and the availability of food reinforcement (e.g., a 50-s fixed interval). During the overall interval a sound stimulus was used to signal shorter intervals that divided the overall interval into equal segments. When there was a fixed temporal relation between the final segment signal and the availability of reinforcement, there was a double-scallop pattern of responding throughout the segmented overall interval; the function relating response rate to time during segment intervals was similar to the function relating response rate to time in unsegmented overall intervals; a change in response rate occurred at the time that a normally presented segment signal was omitted. Taken together, the results indicate that rats timed the overall interval and the segment intervals simultaneously and independently without interference. In Experiment 7 a light stimulus was used on some trials, and a sound stimulus was used on other trials to signal a discrete-trial 50-s peak procedure. When these two signals were presented in compound, there was a leftward shift of the response function, which suggests that rats timed both signals simultaneously. For all of the experiments a scalar timing model with specific stimulus integration rules is used to explain the results. The stimulus integration rule used in the first six experiments, in which there were two signals for the same reinforcement, was to respond if both the segment and the overall interval had exceeded a response threshold. The stimulus integration rule used in Experiment 7, in which there were two signals for different reinforcements, was to respond if the response threshold for either interval had been exceeded.     

Human performance on an analogue of an interval bisection task

Two experiments used normal adult human subjects in an analogue of a time interval bisection task frequently used with animals. All presented durations were defined by the time between two very brief clicks, and all durations were less than 1 sec, to avoid complications arising from chronometric counting. In Experiment 1 different groups of subjects received standard durations of either 0.2 and 0.8 or 0.1 and 0.9 sec and then classified a range of durations including these values in terms of their similarity to the standard short (0.2- or 0.1-sec) and long (0.8- or 0.9-sec) durations. The bisection point (defined as the duration classified as "long" on 50% of trials) was located at 0.43 sec in the 0.2-0.8 group, and at 0.46 sec in the 0.1-0.9 group. Experiment 2 replicated Experiment 1 using a within-subject procedure. The bisection point of both 0.2- and 0.8 sec and 0.1- and 0.9-sec durations was found to be 0.44 sec. Both experiments thus found the bisection point to be located at a duration just lower than the arithmetic mean of the standard short and long durations, rather than at the geometric mean, as in animal experiments. Some other performance measures, such as difference limen, and Weber ratio, were, however, of similar values to those found in bisection tasks with animals. A theoretical model assuming that humans bisect by taking the difference between a presented duration and the short and long standards, as well as having a bias to respond "long", fitted the data well. The model incorporated scalar representations of standard durations and thus illustrated a way in which the obtained results, although different from those found with animal subjects, could be reconciled with scalar timing theory.     

Choice in the time-left procedure and in concurrent chains with a time-left terminal link.

In two experiments, rats chose between a standard fixed-duration food-associated stimulus and a stimulus whose duration was the time remaining to reinforcement in an elapsing comparison interval. In Experiment 1, 4 rats responded in a time-left procedure wherein a single initial-link variable-interval schedule set up two potential terminal links simultaneously. As time elapsed in the initial-link schedule, the choice was between a standard fixed-interval 30-s terminal link and a time-left terminal link whose programmed interval requirement equaled 90 s minus the elapsed time in the initial link. Rats generally responded more on the lever with the shortest programmed terminal-link duration, but the temporal parameters of the procedure were found to vary with response distributions. Contrary to previous reports, therefore, time-left data were well predicted by choice models that make no assumptions about animal timing. In Experiment 2, 8 rats responded on a concurrent-chains schedule with independent variable-interval initial links and a time-left terminal link in one of the choice schedules. On the time-left lever, the programmed terminal-link delay equaled 90 s minus the elapsed time in the time-left initial link. On the standard lever, terminal-link responses were reinforced according to a variable-interval schedule whose average value varied over four conditions. Relative time-left initial-link responses increased in the elapsing time-left initial-link schedule as the time-left terminal link became shorter relative to the standard terminal link. Scalar expectancy theory failed to predict the resultant data, but a modified version of the delay-reduction model made good predictions. An analysis of the elaboration of scalar expectancy theory for variable delays demonstrated that the model is poorly formulated for arithmetically distributed delays.     

Good vibrations: Human interval timing in the vibrotactile modality

This article reports a detailed examination of timing in the vibrotactile modality and comparison with that of visual and auditory modalities. Three experiments investigated human timing in the vibrotactile modality. In Experiment 1, a staircase threshold procedure with a standard duration of 1,000 ms revealed a difference threshold of 160.35 ms for vibrotactile stimuli, which was significantly higher than that for auditory stimuli (103.25 ms) but not significantly lower than that obtained for visual stimuli (196.76 ms). In Experiment 2, verbal estimation revealed a significant slope difference between vibrotactile and auditory timing, but not between vibrotactile and visual timing. That is, both vibrations and lights were judged as shorter than sounds, and this comparative difference was greater at longer durations than at shorter ones. In Experiment 3, performance on a temporal generalization task showed characteristics consistent with the predications of scalar expectancy theory (SET: Gibbon, 1977) with both mean accuracy and scalar variance exhibited. The results were modelled using the modified Church and Gibbon model (MCG; derived by Wearden, 1992, from Church & Gibbon 1982). The model was found to give an excellent fit to the data, and the parameter values obtained were compared with those for visual and auditory temporal generalization. The pattern of results suggest that timing in the vibrotactile modality conforms to SET and that the internal clock speed for vibrotactile stimuli is significantly slower than that for auditory stimuli, which is logically consistent with the significant differences in difference threshold that were obtained.     

Traveling in time: a time-left analogue for humans

Two experiments studied normal humans in an analogue of the time-left procedure of J. Gibbon and R. M. Church (1981). In Experiment 1 the "standard" alternative (S) was always half the length of the "comparison" time-left link (C), and S ranged from 4 to 8 s. Humans showed an increasing preference for the time-left alternative with increasing elapsed time in the interval, and indifference points strongly supported the idea of a linear, rather than a logarithmic, time scale. Experiment 2 used some conditions in which S was greater or less than C/2, and preference for the time-left alternative varied systematically with the S/C ratio. Data from both experiments showed reasonable superposition, suggesting underlying scalar timing processes in time left in humans.     

Sources of variability and systematic error in mouse timing behavior

In the peak procedure, starts and stops in responding bracket the target time at which food is expected. The variability in start and stop times is proportional to the target time (scalar variability), as is the systematic error in the mean center (scalar error). The authors investigated the source of the error and the variability, using head poking in the mouse, with target intervals of 5 s, 15 s, and 45 s, in the standard procedure, and in a variant with 3 different target intervals at 3 different locations in a single trial. The authors conclude that the systematic error is due to the asymmetric location of start and stop decision criteria, and the scalar variability derives primarily from sources other than memory.     

Time perception with and without a concurrent nontemporal task

Prospective time estimates were obtained from human subjects for stimulus durations ranging from 2 to 23 sec. Presence and absence of a concurrent nontemporal task was manipulated within subjects in three experiments. In addition, location of the task within temporal reproduction trials and psychophysical method were varied between groups in Experiments 2 and 3, respectively. For long-duration stimuli, the results of all three experiments conformed to results in the literature, showing a decrease in perceived duration under concurrent task conditions, in accord with attentional resource allocation models of timing. The effects of task location and psychophysical method on time estimates were also compatible with this analysis. However, psychophysical functions obtained under task conditions were fit well by power functions, an outcome that would not be anticipated on the basis of attention theory. The slopes of the functions under no-task conditions were steeper than those under task conditions. The data support the perceptual hypothesis that different sources of sensory input mediate timing under task and no-task conditions.     

Force and timing variability in rhythmic unimanual tapping

In 3 experiments the interdependencies between timing and force production in unimanual paced and self-paced rhythmic tapping tasks were examined as participants (N = 6 in each experiment) tapped (a) to 1 of 3 target periods (333 ms, 500 ms, and 1,000 ms), while they simultaneously produced a constant peak force (PF) over a 50-s trial; (b) to produce 1 of 3 target forces (5, 10, and 15 N) at their preferred frequency, while keeping their rhythm as invariant as possible; and (c) to all combinations of target force and period. The results showed that (a) magnitudes of force and period were largely independent; (b) variability in timing increased proportionally with tapping period, and the variability in force increased with peak force; (c) force variability decreased at faster tapping rates; and (d) timing variability decreased with increasing force levels. (e) Analysis of tap-to-tap variability revealed adjustments over sequences of taps and an acceleration in the tapping rate in unpaced conditions. The interdependencies of force and time are discussed with respect to the challenges they provide for an oscillator-based account.     

Temporal control of conditioned responding in goldfish

The peak procedure was used to characterize response timing during acquisition and maintenance of conditioned responding in goldfish. Subjects received light-shock pairings with a 5- or 15-s interstimulus interval. On interspersed peak trials, the conditioned stimulus light was presented for 45 s and no shock was delivered. Peaks in the conditioned response, general activity, occurred at about the time of the expected unconditioned stimulus, and variability in the activity distribution was scalar. Modeling of the changes in the activity distributions over sessions revealed that the temporal features of the conditioned response changed very little during acquisition. The data suggest that times are learned early in training, and, contrary to I. P. Pavlov's (1927/1960) concept of "inhibition of delay," that timing is learning when to respond rather than learning when not to respond.     

Measuring the Ponzo illusion with the method of production

The Ponzo illusion was varied as a function of angle of oblique arms and was measured by the method of production. The results showed that the method of production produced very similar results to those obtained by the method of limits. The data also indicated that relatively stable individual scores could be obtained if about six judgments were required and if the targets elicited a large illusion. It was concluded that the method of production should be given serious consideration when choosing a psychophysical method.     

Multiple timing and the allocation of attention

Two experiments were designed to examine the effects of multiple timing tasks on prospective time judgment performance. In experiment 1, subjects were required to monitor the durations of one, two, three, or four concurrent target stimuli which began and ended at different times, and then reproduce one of those durations subsequently chosen at random. Time judgment accuracy deteriorated as the number of target stimuli increased. In experiment 2, subjects used the production method to generate specified durations for one, two, three, or four partially overlapping stimuli. Timing was less accurate in conditions involving more target stimuli. In the multiple-target conditions, time judgments were less accurate for the later- rather than earlier-onset targets. The results support an attentional model of timing, and suggest that timing is an effortful process which draws from limited attentional resources.