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The United States armed forces have a lead on a suspected terrorist in the Middle East, but as the man is a recluse, getting a positive ID proves difficult, and the DOD's computer system recommends that the mission be aborted. The Secretary of Defense (Michael Chiklis) agrees with the abort recommendation, but the President orders the mission be carried out anyway. This turns into a political backlash when all those killed turn out to be civilians, and retaliatory bombings are carried out in response.

Jerry Shaw (Shia LaBeouf), is a Stanford University dropout who lacks direction and faces financial difficulty. He finds out his twin brother Ethan, is dead. Following the funeral he goes to withdraw some money from an ATM and is surprised to see that he has $750,000 in his account. When he returns home he finds his apartment filled with a large amount of weapons, explosives, and forged documents. He receives a phone call from an unknown woman, who explains that the FBI is about to apprehend him in thirty seconds and that he must escape. Not believing her, he is caught by the FBI Agent Thomas Morgan (Billy Bob Thornton) and Air Force Office of Special Investigations Special Agent Zoe Perez (Rosario Dawson). The unknown woman arranges Jerry's escape and has him join up with single mother Rachel Holloman (Michelle Monaghan). Holloman is being coerced into helping by the woman, who threatens to kill her son, Sam, a trumpet player on his way to Washington, D.C. from Chicago for a band recital.

The woman helps the pair to avoid the police and FBI units, demonstrating the ability to remotely control virtually any networked device, such as traffic lights, cell phones, and even automated cranes. While Jerry and Rachel follow her instructions, the woman has a crystal explosive made into a necklace and its sound-based trigger placed inside Sam's trumpet. Jerry and Rachel are led from Chicago to Washington, D.C. via Indianapolis and Dayton through various means. Along the way, they are introduced to the woman, who is actually a top secret super-computer called ARIA tasked with gathering intelligence from all over the world. ARIA, the overtaken computer, can control virtually anything electronic to that end. In light of the mistake made by the President at the beginning of the film, ARIA has decided that the executive branch is a threat to the public good and must be eliminated. ARIA plans to destroy the President's cabinet, and calls this Operation Guillotine. It has decided to leave the Secretary of Defense, who agreed with its recommendation to abort, as the successor to the presidency. She does not reveal this to Jerry or Rachel, merely explaining that she is trying to help the people of the United States.

At the Pentagon, where ARIA is housed, Agent Perez discovers that Ethan worked as a technician for the computer and locked it down to prevent ARIA from carrying out her plan. Perez warns the Secretary of Defense and they discuss the situation in a sealed room to prevent ARIA from hearing their conversation. Jerry and Rachel arrive at the Pentagon and are led to the super computer, where ARIA forces Jerry to impersonate Ethan and use an override code allowing her to go ahead with the plan. Jerry observes CCTV footage displaying Ethan's fatal car crash (due to sabotaged traffic lights), realizing ARIA orchestrated Ethan's death since his twin could've stopped her. She then instructs Rachel to eliminate Jerry to prevent the lock from being reinstated, but Rachel cannot bring herself to do it. Rachel is led out of the building by ARIA while Jerry is caught by Agent Morgan. Having been warned by Agent Perez, Morgan believes Jerry's story and takes him to the United States Capitol. On their way, however, ARIA sends an MQ-9 Reaper UCAV after them. Agent Morgan sacrifices himself to destroy the craft and save Jerry.

Meanwhile, Agent Perez returns to the super computer and is successful in destroying it (by piercing her positronic CPU), but not before ARIA uploads 27% of its memory to another location. Rachel is given the explosive necklace and sent to watch the President's speech. Sam's class, whose recital has been moved from the Kennedy Center to the Capitol for the President's State of the Union Address, begins to play. The trigger that will set off the explosive necklace is set to activate when Sam plays a high F on his trumpet corresponding to the word "free" in the last verse of the U.S. national anthem. Jerry successfully infiltrates the vicinity and fires Morgan's pistol into the air, stopping the performance and emptying the room just as Sam starts to play the the F, but provoking a Secret Service agent to shoot him in the shoulder.

In the aftermath of the chaos ARIA caused, the Secretary of Defense urges that another super computer should not be built. Ethan posthumously receives the Medal of Honor while Jerry, injured but alive and well, receives the Congressional Gold Medal. The film ends with Jerry attending Sam's birthday party. Jerry gives Sam the video game Rock Band and Sam and his friends go off and play it. Rachel thanks him for attending and kisses him on the cheek. She then tells Jerry that she is glad that he is there.