Mirror neurons within a monkey's premotor area F5 fire not only when the monkey performs a certain class of action but also when the monkey observes another monkey (or the experimenter) perform a similar action (Gallese et al. 1996;Rizzolatti et al. 1996a) . It has thus been argued that these neurons are crucial for understanding of actions by others. We offer the Hand-State Hypothesis as a new explanation of the evolution of this capability, hypothesizing that these neurons first evolved to augment the "canonical" F5 neurons (active during self-movement based on observation of an object) by providing visual feedback on "hand state", relating the shape of the hand to the shape of the object. We then introduce the MNS (Mirror Neuron System) model of F5 and related brain regions. The existing FARS (Fagg-Arbib-Rizzolatti-Sakata) model (Fagg and Arbib 1998) represents circuitry for visually-guided grasping of objects, linking parietal area AIP with F5 canonical neurons. The MNS model extends the AIP visual pathway by also modeling pathways, directed toward F5 mirror neurons, which match arm-hand trajectories to the affordances and location of a potential target object. We present the basic schemas for the MNS model, then aggregate them into three "grand schemas" − Visual Analysis of Hand State, Reach and Grasp, and the Core Mirror Circuit − for each of which we present a useful implementation. With this implementation we show how the mirror system may learn to recognize actions already in the repertoire of the F5 canonical neurons. We show that the connectivity pattern of mirror neuron circuitry can be established through training, and that the resultant network can exhibit a range of novel, physiologically interesting, behaviors during the process of action recognition. We train the system on the basis of final grasp but then observe the whole time course of mirror neuron activity, yielding predictions for neurophysiological experiments under conditions of spatial perturbation, altered kinematics, and ambiguous grasp execution which highlight the importance of the timing of mirror neuron activity.