Why it is a question
THEORY-909 holds that, in five papers reviewed by Gerstner et al., mostly slice experiments with one laboratory per region, a pairing changed the synapse when a neuromodulator or plateau potential followed within one to five seconds and not later. The reviewers grant that persistent network activity or short-term plasticity could bridge the same gap. CLAIM-443’s schematic three-factor rule would need such a trace to learn from delayed outcomes, though it claims no neural implementation.
What would answer it
The seconds-scale trace shown in behaving animals, not only in slices: an experiment in vivo in which interfering with the trace’s mechanism (for example CaMKII in striatal spines) during the interval between action and delayed outcome abolishes learning from that outcome while leaving learning from immediate outcomes intact, and independent replication of at least the striatal and cortical time constants. Refuted if learning from delays of seconds survives the abolition of synaptic traces, carried instead by persistent network activity or short-term plasticity. Further slice experiments with the same protocols cannot settle it.