A popular view of interval timing in animals is that it is driven by a discrete pacemaker-accumulator mechanism that yields a linear scale for encoded time. But these mechanisms are fundamentally at odds with the Weber law property of interval timing, and experiments that support linear encoded time can be interpreted in other ways. We argue that the dominant pacemaker-accumulator theory, scalar expectancy theory (SET), fails to explain some basic properties of operant behavior on interval-timing procedures and can only accommodate a number of discrepancies by modifications and elaborations that raise questions about the entire theory. We propose an alternative that is based on principles of memory dynamics derived from the multiple-time-scale (MTS) model of habituation. The MTS timing model can account for data from a wide variety of time-related experiments: proportional and Weber law temporal discrimination, transient as well as persistent effects of reinforcement omission and reinforcement magnitude, bisection, the discrimination of relative as well as absolute duration, and the choose-short effect and its analogue in number-discrimination experiments. Resemblances between timing and counting are an automatic consequence of the model. We also argue that the transient and persistent effects of drugs on time estimates can be interpreted as well within MTS theory as in SET. Recent real-time physiological data conform in surprising detail to the assumptions of the MTS habituation model. Comparisons between the two views suggest a number of novel experiments.
Pigeons tracked sinusoidal sequences of interfood intervals (IFIs) by pausing in each interval for a time proportional to the preceding interval. Schedules with either long (30-90 s) or short (5-15 s) values, with variable numbers of cycles and starting phase each day, were tracked about equally well. Tracking was apparently immediate and did not improve across sessions. Experiment 2, in which long and short series were presented on alternate days, showed that tracking on long was more impaired than on short. Experiment 3 showed that occasional presentation of a short IFI in a series of fixed, longer IFIs caused a reduction in waiting time in the next IFI. These effects are evidence for a fast-acting timing mechanism in which waiting time in the IFI N + 1 is strongly determined by the preceding IFI, N. Earlier IFIs have some cumulative effect, but the details remain to be elucidated.
Habituation is the waning of a reflex response to repeated stimulation. Habituation to closely spaced stimuli is faster and more complete than to widely spaced stimuli, but recovery is also more rapid (rate sensitivity). We show that a 2-unit, cascaded-integrator dynamic model can explain in detail an extensive data set on rate-sensitive habituation in the nematode Caenorhabditis elegans. Many apparently complex properties of habituation and learning dynamics may reflect interactions among a small number of processes with different time scales.
Recent developments reveal that animals can rapidly learn about intervals of time. We studied the nature of this fast-acting process in two experiments. In Experiment 1 pigeons were exposed to a modified fixed-time schedule, in which the time between food rewards (interfood interval) changed at an unpredictable point in each session, either decreasing from 15 to 5 s (step-down) or increasing from 15 to 45 s (step-up). The birds were able to track under both conditions by producing postreinforcement wait times proportional to the preceding interfood-interval duration. However, the time course of responding differed: Tracking was apparently more gradual in the step-up condition. Experiment 2 studied the effect of having both kinds of transitions within the same session by exposing pigeons to a repeating (cyclic) sequence of the interfood-interval values used in Experiment 1. Pigeons detected changes in the input sequence of interfood intervals, but only for a few sessions-discrimination worsened with further training. The dynamic effects we observed do not support a linear waiting process of time discrimination, but instead point to a timing mechanism based on the frequency and recency of prior interfood intervals and not the preceding interfood interval alone.
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