An Analysis of the Brain Systems Involved with Producing Letters by Hand.

Complex visual-motor behaviors dominate human-environment interactions. Letter production, writing individual letters by hand, is an example of a complex visual-motor behavior composed of numerous behavioral components, including the required motor movements and the percepts that those motor movements create. By manipulating and isolating components of letter production, we provide experimental evidence that this complex visual-motor behavior is supported by a widespread neural system that is composed of smaller subsystems related to different sensorimotor components. Adult participants hand-printed letters with and without "ink" on an MR-safe digital writing tablet, perceived static and dynamic representations of their own handwritten letters, and perceived typeface letters during fMRI scanning. Our results can be summarized by three main findings: (1) Frontoparietal systems were associated with the motor component of letter production, whereas temporo-parietal systems were more associated with the visual component. (2) The more anterior regions of the left intraparietal sulcus were more associated with the motor component, whereas the more posterior regions were more associated with the visual component, with an area of visual-motor overlap in the posterior intraparietal sulcus. (3) The left posterior intraparietal sulcus and right fusiform gyrus responded similarly to both visual and motor components, and both regions also responded more during the perception of one's own handwritten letters compared with perceiving typed letters. These findings suggest that the neural systems recruited during complex visual-motor behaviors are composed of a set of interrelated sensorimotor subsystems that support the full behavior in different ways and, furthermore, that some of these subsystems can be rerecruited during passive perception in the absence of the full visual-motor behavior.