Delay-dependent transitions of phase synchronization and coupling symmetry between neurons shaped by spike-timing-dependent plasticity

Synchronization plays a key role in learning and memory by facilitating the communication between neurons promoted by synaptic plasticity. Spike-timing-dependent plasticity (STDP) is a form of synaptic plasticity that modifies the strength of synaptic connections between neurons based on the coincidence of pre- and postsynaptic spikes. In this way, STDP simultaneously shapes the neuronal activity and synaptic connectivity in a feedback loop. However, transmission delays due to the physical distance between neurons affect neuronal synchronization and the symmetry of synaptic coupling. To address the question that how transmission delays and STDP can jointly determine the emergent pairwise activity-connectivity patterns, we studied phase synchronization properties and coupling symmetry between two bidirectionally coupled neurons using both phase oscillator and conductance-based neuron models. We show that depending on the range of transmission delays, the activity of the two-neuron motif can achieve an in-phase/anti-phase synchronized state and its connectivity can attain a symmetric/asymmetric coupling regime. The coevolutionary dynamics of the neuronal system and the synaptic weights due to STDP stabilizes the motif in either one of these states by transitions between in-phase/anti-phase synchronization states and symmetric/asymmetric coupling regimes at particular transmission delays. These transitions crucially depend on the phase response curve (PRC) of the neurons, but they are relatively robust to the heterogeneity of transmission delays and potentiation-depression imbalance of the STDP profile.