Both animals and humans are capable of representing and comparing numerical quantities, but only humans seem to have evolved multi-digit place-value number systems. This article develops a neural model, called the Spatial Number Network, or SpaN model, which predicts how these shared numerical capabilities are computed using a spatial representation of number quantities in the Where cortical processing stream, notably the Inferior Parietal Cortex. Multi-digit numerical representations that obey a place-value principle are proposed to arise through learned interactions between categorical language representations in the What cortical processing stream and the Where spatial representation. It is proposed that learned semantic categories that symbolize separate digits, as well as place markers like "tens," "hundreds," "thousands," etc., are associated through learning with the corresponding spatial locations of the Where representation, leading to a place-value number system as an emergent property of What-Where information fusion. The model quantitatively simulates error rates in quantification and numerical comparison tasks, and reaction times for number priming and numerical assessment and comparison tasks. In the Where cortical process, it is proposed that transient responses to inputs are integrated before they activate an ordered spatial map that selectively responds to the number of events in a sequence. Neural mechanisms are defined which give rise to an ordered spatial numerical map ordering and Weber law characteristics as emergent properties. The dynamics of numerical comparison are encoded in activity pattern changes within this spatial map. Such changes cause a "directional comparison wave" whose properties mimic data about numerical comparison. These model mechanisms are variants of neural mechanisms that have elsewhere been used to explain data about motion perception, attention shifts, and target tracking. Thus, the present model suggests how numerical representations may have emerged as specializations of more primitive mechanisms in the cortical Where processing stream. The model's What-Where interactions can explain human psychophysical data, such as error rates and reaction times, about multi-digit (base 10) numerical stimuli, and describe how such a competence can develop through learning. The SpaN model and its explanatory range arc compared with other models of numerical representation.Defense Advanced Research Projects Agency and the Office of Naval Research (N00014-95-1-0409); National Science Foundation (IRI-97-20333
