Stages of nonsymbolic number processing in occipitoparietal cortex disentangled by fMRI adaptation

The neurobiological mechanisms of nonsymbolic number processing in humans are still unclear. Computational modeling proposed three successive stages: first, the spatial location of objects is stored in an object location map; second, this information is transformed into a numerical summation code; third, this summation code is transformed to a number-selective code. Here, we used fMRI-adaptation to identify these three stages and their relative anatomical location. By presenting the same number of dots on the same locations in the visual field, we adapted neurons of human volunteers. Occasionally, deviants with the same number of dots at different locations or different numbers of dots at the same location were shown. By orthogonal number and location factors in the deviants, we were able to calculate three independent contrasts, each sensitive to one of the stages. We found an occipitoparietal gradient for nonsymbolic number processing: the activation of the object location map was found in the inferior occipital gyrus. The summation coding map exhibited a nonlinear pattern of activation, with first increasing and then decreasing activation, and most activity in the middle occipital gyrus. Finally, the number-selective code became more pronounced in the superior parietal lobe. In summary, we disentangled the three stages of nonsymbolic number processing predicted by computational modeling and demonstrated that they constitute a pathway along the occipitoparietal processing stream.