Interval Timing
Time Scales of Behavioral Timing
Researchers separate timing into three distinct scales, each with unique behavioral signatures, neural circuitry and plasticity mechanisms.
• Circadian (≈ 24 h)
Governs sleep–wake cycles, daily appetite fluctuations, arousal patterns.
Key structure: in the hypothalamus.
SCN lesions → fragmented sleep (many short naps throughout the day).
Mechanism: transcription/translation feedback loops in clock-genes; selective breeding can shorten or lengthen rhythms ⇒ strong genetic component.
Precision: high.
Flexibility/adaptability: low (slow to shift).
• Interval (≈ seconds → minutes) – principal focus of the lecture.
Supports decision making at traffic lights, elevator waiting, planning coffee runs, animal foraging (e.g., bees revisiting a flower after nectar replenishment).
Brain circuits: meso-cortico-striatal dopaminergic loops (striatum ⇄ frontal cortex).
Mechanism of adjustment: synaptic plasticity (e.g., ) rather than gene transcription.
Precision: moderate (estimates contain noise).
Flexibility: high (rapid learning & updating).
• Millisecond
Underlies rapid, fine-grained motor acts: musical performance, speech articulation, 3-point basketball shots.
Cerebellum heavily implicated; may also rely on intrinsic cellular oscillations + LTP.
Precision: very high.
Flexibility: also high ("best of both worlds").
Interval Timing & Operant Schedules
Historical roots: analysis of reinforcement schedules.
Four classic instrumental schedules reviewed:
• (Variable Ratio)
• (Fixed Ratio) – produces run–break pattern; possibly a metronomic effect rather than literal counting.
• (Variable Interval) – steady, low response rate, minimal post-reinforcement pause.
• (Fixed Interval) – produces "scalloping": slow start → accelerating responses → abrupt stop at reinforcement.Key experiment (Roberts 1981 – Phase 1)
• Light → FI20 Tone → FI40
• On 20 % of trials, food omitted ("empty" trials) to reveal internal clock independent of reinforcement.
• Response-rate curves peaked at ≈ 20 s (light) or 40 s (tone).
• Wider generalization gradient at 40 s ⇒ lower precision for longer intervals.
Dissociating Motivation and Timing (Roberts 1981 – Phase 2)
Both stimuli now but probability of reinforcement manipulated:
• Light: food on 80 % of trials.
• Tone: food on 20 % of trials.Findings:
• Peak response time remained ≈ 20 s for both stimuli (unchanged clock).
• Peak response rate differed dramatically (higher under 80 % chance) ⇒ motivation affects rate, not timing perse.Conclusion: temporal control and motivational variables can be experimentally dissociated.
Human Parallels in Timing Variability (Ratkin 1998)
Human subjects view a target duration (, or s), then hold a button and release when they believe the same duration has elapsed.
Resulting release-probability curves mirror rat data:
• Narrow, high-precision peaks for short durations.
• Broader, noisier distributions as duration lengthens.Demonstrates cross-species commonality in scalar property of timing.
Direction of the Internal Clock: Count-Up vs Count-Down
Test paradigm (Roberts 1981)
• Training: Tone → FI20 → LightFI40
• Probe: Present tone for , or s, then switch to light.Predictions:
• Countdown model – peak should occur at (e.g., 5 s tone ⇒ 35 s peak).
• Count-up model – peak should stay at irrespective of pre-light tone length.Data: All three conditions peaked at ⇒ animals use stopwatch-like count-up mechanism.
Ability to Pause and Resume the Clock
Timeout experiment (Roberts 1981)
• Baseline: Tone → (animals well-trained after ≈ 19 days).
• Test: During empty trial, lever retracted & house light off for (or ).Results:
• Peak response shifted rightward by ≈ timeout length (10 s blackout → 50 s peak; 5 s → 45 s peak).
• Peak response rate unaffected ⇒ motivation stable.Interpretation: Animals can stop accumulating time during blackout and restart at prior value.
Scalar Timing Theory (Gibbon 1977)
Components (functional boxes):
Pacemaker – tonic pulse generator ("metronome").
Switch – closes when timing stimulus appears, routing pulses to…
Accumulator (Working Memory) – counts pulses; represents current elapsed time.
Reference Memory – stores mean pulse count for past reinforced intervals under that stimulus.
Comparator / Decision Rule – continually checks similarity between accumulator and reference; greater similarity ⇒ higher response probability.
Explains:
• Count-up direction (switch starts at 0 pulses).
• Pause-resume (opening switch freezes count).
• Scalar property (constant coefficient of variation when pulses follow Poisson process).
Neurobiological Substrates & Pharmacology
Pacemaker speed modulated by dopamine in cortico-striatal loops.
Methamphetamine (dopamine agonist) – accelerates pacemaker.
• Initial effect: peak times earlier (e.g., FI-40 peak shifts to ≈ 30 s; FI-20 to ≈ 10 s).
• With continued drug exposure, reference memory recalibrates ⇒ peaks drift back toward nominal interval.
• After drug removal (saline), pacemaker slows to baseline but reference memory "expects" more pulses ⇒ peaks occur later (≈ 50 s & 30 s).Haloperidol (dopamine antagonist) – produces mirror-image pattern (slowed pacemaker, delayed peaks, then opposite rebound).
Supports link between dopamine tone and clock speed.
Practical, Philosophical & Ethical Implications
Real-world relevance: understanding drug-induced distortions of time (amphetamine, antipsychotics) in humans.
Motor-skill coaching: millisecond cerebellar timing insights help optimize athletic & musical training.
Animal-welfare & research ethics: recognizing precise vs flexible timing capacities informs enrichment schedules and experimental design.
Clinical translation: circadian-striatum-cerebellum distinctions guide treatment of sleep disorders, ADHD, Parkinson’s (dopaminergic timing deficits).