We all have a sense of time. Yet, there are no sensory receptors specifically dedicated for perceiving time. It is an almost uniquely intangible sensation: we cannot see time in the way that we see color, shape, or even location. So how is time represented in the brain? We explore the neural substrates of metrical representations of time such as duration estimation (explicit timing) or temporal expectation (implicit timing). Basal ganglia (BG), supplementary motor area, cerebellum, and prefrontal cortex have all been linked to the explicit estimation of duration. However, each region may have a functionally discrete role and will be differentially implicated depending upon task context. Among these, the dorsal striatum of the BG and, more specifically, its ascending nigrostriatal dopaminergic pathway seems to be the most crucial of these regions, as shown by converging functional neuroimaging, neuropsychological, and psychopharmacological investigations in humans, as well as lesion and pharmacological studies in animals. Moreover, neuronal firing rates in both striatal and interconnected frontal areas vary as a function of duration, suggesting a neurophysiological mechanism for the representation of time in the brain, with the excitatory-inhibitory balance of interactions among distinct subtypes of striatal neuron serving to fine-tune temporal accuracy and precision.

Attention modulates our subjective perception of time. The less we attend to an event's duration, the shorter it seems to last. Attention to time or color stimulus attributes was modulated parametrically in an event-related functional magnetic resonance imaging study. Linear increases in task performance were accompanied by corresponding increases in brain activity. Increasing attention to time selectively increased activity in a corticostriatal network, including pre-supplementary motor area and right frontal operculum. Increasing attention to color selectively increased activity in area V4. By identifying areas whose activity was specifically modulated by attention to time, we have defined the core neuroanatomical substrates of timing behavior.

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