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The movie opens with a TV show featuring Chef Auguste Gusteau (Brad Garrett), owner of the best restaurant in Paris, talking about his bestselling cookbook, which proudly bears his mantra "Anyone Can Cook!"

A rat named Remy (voice of Patton Oswalt) begins talking about his life in monologue fashion. Remy states that he has enhanced senses of both taste and smell, which makes him very particular about what he eats. Remy's brother Emile (Peter Sohn) is impressed by this talent, but their father Django (Brian Dennehy) who leads the rats' colony, could care less - until Remy reveals that he can recognize the scent of rat poison in or near food. Django puts Remy to work sniffing and testing food for the rest of the clan. Remy is not happy about the rats having to steal food from the garbage; he would prefer to go to the kitchen and take the "fresh" samples. But Django hates and fears humans so he forbids Remy (and all other members of the clan) to interact with them.

Despite his father's orders, Remy spends several nights in a human home, reading Chef Gusteau's cookbook and watching television programs about cooking. Before long he has a near-expert level of knowledge about food preparation. One day, Remy takes Emile into the kitchen to get some spices that will go with some other food samples they have gathered. Emile hesitates, but agrees to go with his brother. While inside, Remy sees on TV that a famous food critic named Anton Ego (Peter O'Toole), gave Gusteau's restaurant a less-than-stellar review that resulted in the restaurant losing one of its five stars. A heartbroken Gusteau died soon after, which meant the loss of another star according to tradition.