Giselle lives in the blissful and traditionally-animated world of Andalasia, where animals are talkative companions and musical interludes punctuate nearly every interaction. She dreams of her true love and as she sings about True Love's Kiss to her chipmunk friend Pip and other forest animals, Prince Edward hears her voice in the forest. After he rescues her from a troll, they decide to marry the next day. However, Edward's step-mother, Queen Narissa, will lose her claim to the throne upon Edward's marriage, so when Giselle arrives at the palace, Narissa disguises herself as an old hag and throws Giselle down a well and into a magic portal to a world "where there are no happily ever afters" in order to keep her step-son single.
Giselle emerges through a manhole in Times Square, in modern-day, live-action New York City, and after a turn of events meets Robert Philip, a cynical, realistic divorce lawyer who is at first reluctant to help her find her way home. He allows her to stay at his apartment despite believing that she is crazy and worrying about the safety of his young daughter Morgan . To Robert's surprise, Giselle invites animals in the city?rats, pigeons and cockroaches?to help her clean up his apartment, and constructs a dress using material cut from his curtains. Robert reaches the end of his patience when Giselle causes an argument between him and his soon-to-be-fiancée Nancy, and a scene at his office because of her naivete.
He decides to part with Giselle at Central Park, but rejoins her after seeing her give the money he gave her to an old woman. During their walk through Central Park, Giselle questions Robert on how he displays his affection for Nancy and spontaneously starts the musical production number "That's How You Know" with many performers in the park joining her. Giselle helps Robert reconcile with Nancy by sending an apology on Robert's behalf, along with tickets to the King's and Queen's Ball.
Meanwhile, Queen Narissa's henchman Nathaniel follows Edward and Pip, who have journeyed to New York to save Giselle. They stop at a motel, where Nathaniel questions himself and his relationship with Narissa after watching a soap opera. He sneaks out to give Giselle a poisoned apple. Pip is unable to speak coherently in this world and has a frustrating time alerting the Prince of the minion's intentions. When Nathaniel fails twice to poison Giselle, Narissa becomes infuriated.
As Giselle and Robert spend more time together, Giselle discovers that the real world is much more complicated than she realized, while Robert is affected by her optimism and idealism. Edward continues to look for Giselle and eventually finds her at Robert's apartment. While Edward is eager to take Giselle home, they go on a date around New York at her insistence. To Nancy's chagrin, Giselle and Edward attend the King's and Queen's Ball. After Nancy and Edward pair off to dance, Giselle dances with Robert. During their dance, Giselle realizes that Robert is her true love. Unbeknownst to them, Narissa has traveled to New York from Andalasia. Under her old hag's disguise, she manages to successfully poison Giselle, but is stopped by Edward before she can escape with Giselle's unconscious form.
A remorseful Nathaniel reveals Narissa's plot, admits his deeds and reveals that the spell of the poison apple has to be broken before midnight, or Giselle will die. Robert revives Giselle with a true love's kiss, but Narissa uses the distracting moment to break free. She transforms into a dragon, and decides to write her own ending to the story. Taking Robert hostage, she lures Giselle out the window and up to the top of the Woolworth Building. With help from Pip, Narissa falls from the roof to her death, exploding into magic dust at street level. Giselle catches Robert, and they manage to keep themselves from falling off the roof.
Nancy leaves with Edward to Andalasia and marries him. Giselle opens a boutique in New York City, where she is assisted by both humans and animals. Both Nathaniel and Pip become successful authors. The last scene shows Giselle, Robert, and Morgan playing together and living happily ever after as a family.
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