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: suprachiasmatic nucleus (SCN)\text{suprachiasmatic\ nucleus (SCN)} 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., LTP\text{LTP}) 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:
    VR\text{VR} (Variable Ratio)
    FR\text{FR} (Fixed Ratio) – produces run–break pattern; possibly a metronomic effect rather than literal counting.
    VI\text{VI} (Variable Interval) – steady, low response rate, minimal post-reinforcement pause.
    FI\text{FI} (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 FI20s\text{FI}_{20\,\text{s}} 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 (66, 1212 or 2121 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 55, 1010 or 1515 s, then switch to light.

  • Predictions:
    Countdown model – peak should occur at 40tone duration40-\text{tone\ duration} (e.g., 5 s tone ⇒ 35 s peak).
    Count-up model – peak should stay at 40s40\,\text{s} irrespective of pre-light tone length.

  • Data: All three conditions peaked at 40s\approx40\,\text{s} ⇒ animals use stopwatch-like count-up mechanism.

Ability to Pause and Resume the Clock

  • Timeout experiment (Roberts 1981)
    • Baseline: Tone → FI40\text{FI}_{40} (animals well-trained after ≈ 19 days).
    • Test: During empty trial, lever retracted & house light off for 10s10\,\text{s} (or 5s5\,\text{s}).

  • 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):

    1. Pacemaker – tonic pulse generator ("metronome").

    2. Switch – closes when timing stimulus appears, routing pulses to…

    3. Accumulator (Working Memory) – counts pulses; represents current elapsed time.

    4. Reference Memory – stores mean pulse count for past reinforced intervals under that stimulus.

    5. 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).