Getting the timing right: experimental protocols for investigating time with functional neuroimaging and psychopharmacology.

Functional Magnetic Resonance Imaging (fMRI) is an effective tool for identifying brain areas and networks implicated in human timing. But fMRI is not just a phrenological tool: by careful design, fMRI can be used to disentangle discrete components of a timing task and control for the underlying cognitive processes (e.g. sustained attention and WM updating) that are critical for estimating stimulus duration in the range of hundreds of milliseconds to seconds. Moreover, the use of parametric designs and correlational analyses allows us to better understand not just where, but also how, the brain processes temporal information. In addition, by combining fMRI with psychopharmacological manipulation, we can begin to uncover the complex relationship between cognition, neurochemistry and anatomy in the healthy human brain. This chapter provides an overview of some of the key findings in the functional imaging literature of both duration estimation and temporal prediction, and outlines techniques that can be used to allow timing-related activations to be interpreted more unambiguously. In our own studies, we have found that estimating event duration, whether that estimate is provided by a motor response or a perceptual discrimination, typically recruits basal ganglia, SMA and right inferior frontal cortex, and can be modulated by dopaminergic activity in these areas. By contrast, orienting attention to predictable moments in time in order to optimize behaviour, whether that is to speed motor responding or improve perceptual accuracy, recruits left inferior parietal cortex.