Impact of axonal delay on structure development in a multi-layered network

The mechanisms underlying how activity in the visual pathway may give rise through neural plasticity to many of the features observed experimentally in the early stages of visual processing was provided by Linkser in a seminal, three-paper series. Owing to the complexity of multi-layer models, an implicit assumption in Linsker's and subsequent papers has been that propagation delay is homogeneous and plays little functional role in neural behaviour. We relax this assumption to examine the impact of distance-dependent axonal propagation delay on neural learning. We show that propagation delay induces low-pass filtering by dispersing the arrival times of spikes from presynaptic neurons, providing a natural correlation cancellation mechanism for distal connections. The cut-off frequency decreases as the radial propagation delay within a layer increases relative to propagation delay between the layers, introducing an upper limit on temporal resolution. Given that the PSP also acts as a low-pass filter, we show that the effective time constant of each should enable the processing of similar scales of temporal information. This result has implications for the visual system, in which receptive field size and, thus, radial propagation delay, increases with eccentricity. Furthermore, the network response is frequency dependent since higher frequencies require increased input amplitude to compensate for attenuation. This concords with frequency-dependent contrast sensitivity in the visual system, which changes with eccentricity and receptive field size. We further show that the proportion of inhibition relative to excitation is larger where radial propagation delay is long relative to inter-laminar propagation delay. We show that the addition of propagation delay reduces the range in the cell's on-center size, providing stability to variations in homeostatic parameters.