Convergence between cells which differ in both spatial and temporal properties create higher order neurons with response properties that are distinctly different from those of the input neurons. The spatial properties of target neurons are not necessarily "cosinetuned". In addition, unlike the independence between spatial and temporal properties in cosine-tuned afferent neurons, higher-order target cells generally exhibit a dependence of temporal dynamics on spatial properties. The response properties of target neurons receiving spatio-temporal convergence (STC) from tonic and phasic-tonic or phasic afferents is investigated here by considering a general case where the dynamic input is represented by a fractional, leaky, derivative transfer function. It is shown that, at frequencies below the corner frequency of the dynamic input, the temporal properties of target neurons can be described by leaky differentiators having time constants that are a function of spatial direction. Thus, STC target neurons exhibit tonic temporal response properties during stimulation along some spatial directions (having small time constants) and phasic properties along other directions (having large time constants). Specifically, target neurons encode the complete derivative of the stimulus along certain spatial directions. Thus, STC acts as a directionally specific high-pass filter and produces complete derivatives from fractional, leaky derivative afferent signals. In addition, spatio-temporal transformations can generate novel temporal dynamics in the central nervous system. These observations suggest that spatio-temporal computations might constitute an alternative to parallel, independent spatial and temporal channels.
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