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Neural and Behavioral Mechanisms of Interval Timing in the Striatum
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To guide behavior and learn from its consequences, the brain must represent
time over many scales. Yet, the neural signals used to encode time in the
seconds to minute range are not known. The striatum is the major input area of
the basal ganglia; it plays important roles in learning, motor function and
normal timing behavior in the range of seconds to minutes. We investigated
how striatal population activity might encode time. To do so, we recorded the
electrical activity from striatal neurons in rats performing the serial fixed interval
task, a dynamic version of the fixed Interval schedule of reinforcement. The
animals performed in conformity with proportional timing, but did not strictly
conform to scalar timing predictions, which might reflect a parallel strategy to
optimize the adaptation to changes in temporal contingencies and
consequently to improve reward rate over the session. Regarding the neural
activity, we found that neurons fired at delays spanning tens of seconds and
that this pattern of responding reflected the interaction between time and the
animals’ ongoing sensorimotor state. Surprisingly, cells rescaled responses in
time when intervals changed, indicating that striatal populations encoded
relative time. Moreover, time estimates decoded from activity predicted trial-bytrial
timing behavior as animals adjusted to new intervals, and disrupting
striatal function with local infusion of muscimol led to a decrease in timing
performance. Because of practical limitations in testing for sufficiency a
biological system, we ran a simple simulation of the task; we have shown that
neural responses similar to those we observe are conceptually sufficient to
produce temporally adaptive behavior. Furthermore, we attempted to explain
temporal processes on the basis of ongoing behavior by decoding temporal
estimates from high-speed videos of the animals performing the task; we could
not explain the temporal report solely on basis of ongoing behavior. These
results suggest that striatal activity forms a scalable population firing rate code
for time, providing timing signals that animals use to guide their actions.
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Neuroscience