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A girl named Penny and her dog Bolt star on a hit television series Bolt in which the titular character has various superpowers and must constantly thwart the evil plans of the nefarious Dr. Calico. Because Bolt has lived his entire life on a soundstage, and believes he really is his fictional counterpart, he doesn't know how to live like a normal dog. After filming completes for one episode, Bolt escapes from his on-set trailer mistakenly believing Penny has been kidnapped by the television villain. He attempts to break through a window, knocking himself unconscious as he falls into a box of Styrofoam. With no one aware Bolt is in the box, it is shipped from Hollywood to New York. In New York, he meets Mittens, a female alleycat who bullies pigeons out of their food. Because Mittens is a cat similar to one owned by the Green-Eyed Man, Bolt forces her to help him get back to Hollywood, and the two start their journey westward. Meanwhile, in Hollywood, Penny is deeply saddened over Bolt's disappearance but is forced by the studio to continue filming with a Bolt lookalike. Along the way, Bolt starts to notice that his regular superpowers aren't working, but he believes this simply the temporary effect that Styrofoam has on his body, much like Kryptonite affects Superman.

Surprised at his first feelings of both pain and hunger, Bolt manages to get food on the road thanks to the training of Mittens, who shows him how to look like a cute, needy dog. At a campsite, they meet Rhino, a fearless, TV-obsessed hamster and huge Bolt fan who joins their team. Mittens tries to convince Bolt that his superpowers aren't real, but their discussion is cut short by the arrival of Animal Control, who captures them both. Bolt vows to rescue Mittens, and does so with the help of Rhino and an exploding oxygen tank. The cross-country journey continues and Bolt learns to enjoy typical dog activities, (such as hanging his head out the window), but Mittens refuses to go farther than Las Vegas. She tells Bolt that his Hollywood life is fake and there is no real love for him there. Her emotional rant reveals that she was once a housecat, but was abandoned by her previous owner and left to brave the harsh streets alone and declawed. Bolt refuses to believe that Penny doesn't love him, and continues on alone, wishing Mittens the best. Rhino convinces Mittens that they must help him, and the two set off to find Bolt once again.

Bolt returns to the studio and is heartbroken to find Penny embracing his lookalike, unaware that Penny still misses him and that her affection for the lookalike was part of a rehearsal for the show. Witnessing this, and realizing that Penny truly does love Bolt, Mittens confronts Bolt and manages to clear up the misunderstanding. Back on the set, the Bolt-lookalike panics during filming and accidentally knocks over some torches, setting the soundstage on fire and trapping Penny. Bolt arrives and reunites with Penny, and manages to get themselves rescued. Penny's mom has her quit the show because she's has had enough of her agent caring about nothing but business and money. The show continues on with a replacement "Bolt" and "Penny", but seems to have jumped the shark with an alien abduction storyline that even Rhino considers silly. Penny herself adopts Mittens and Rhino, and moves to a rural home to enjoy a simpler, happy lifestyle with Bolt and her new pets.