Probing Our Built-in Calculator: A Systematic Narrative Review of Noninvasive Brain Stimulation Studies on Arithmetic Operation-Related Brain Areas

This systematic review presented a comprehensive survey of studies that applied transcranial magnetic stimulation and transcranial electrical stimulation to parietal and nonparietal areas to examine the neural basis of symbolic arithmetic processing. All findings were compiled with regard to the three assumptions of the triple-code model (TCM) of number processing. Thirty-seven eligible manuscripts were identified for review (33 with healthy participants and 4 with patients). Their results are broadly consistent with the first assumption of the TCM that intraparietal sulcus both hold a magnitude code and engage in operations requiring numerical manipulations such as subtraction. However, largely heterogeneous results conflicted with the second assumption of the TCM that the left angular gyrus subserves arithmetic fact retrieval, such as the retrieval of rote-learned multiplication results. Support is also limited for the third assumption of the TCM, namely, that the posterior superior parietal lobule engages in spatial operations on the mental number line. Furthermore, results from the stimulation of brain areas outside of those postulated by the TCM show that the bilateral supramarginal gyrus is involved in online calculation and retrieval, the left temporal cortex in retrieval, and the bilateral dorsolateral prefrontal cortex and cerebellum in online calculation of cognitively demanding arithmetic problems. The overall results indicate that multiple cortical areas subserve arithmetic skills.