Since winning the Wings Around the Globe race in the Planes (film), Dusty Crophopper has had a successful career as a racer. Unfortunately, his engine's Transmission (mechanics) becomes damaged because Dusty routinely operates the engine beyond its design limits. With that particular model of gearbox now out of production and none available anywhere, Dusty's mechanic Dottie fits a warning light to his control panel to ensure he doesn't damage his gearbox any further. No longer able to race and faced with the possibility of returning to his old job as a crop-duster, Dusty goes on a defiant flight and tests his limits. In doing so, Dusty exceeds his limits and makes a forced landing at Propwash Junction airport, causing a fire.

Dusty Crophopper is a Aerial application plane who works at a cornfield and practices aerobatic maneuvers in his spare time, dreaming of becoming a racer. His dreams are scorned by his boss, Leadbottom, and his Forklift truck/mechanic friend, Dottie. However, he is supported by his fuel truck friend, Chug. Dusty and Chug train for qualifiers for the upcoming Wings Across the Globe race. On the night before the qualifiers, Dusty asks an elderly navy war plane named Skipper Riley to teach him how to fly well, but Skipper refuses. Dusty barely qualifies for the race.

Gregorio and Ingrid Cortez are rival spies who fall in love. They retire and have two children, 12-year-old Carmen and 9-year-old Juni. They work for the Organization of Super Spies (OSS) doing desk work. The children have no idea of their parents' previous career. Gregorio and Ingrid are called back into the field when agents go missing. The children are left in the care of their uncle Felix. Gregorio suspects that children's television host Fegan Floop has kidnapped the agents and mutated them into his "Fooglies," creatures on his show.