In the near future, weapons expert James McCullen (Christopher Eccleston) has created a nanotechnology-based weapon capable of destroying an entire city. His company M.A.R.S. sells four warheads to NATO, and NATO troops are tasked with delivering the warheads. Duke (Channing Tatum) and Ripcord (Marlon Wayans) are delivering the warheads when they are ambushed by the Baroness (Sienna Miller), whom Duke recognizes to be his ex-fiancee Ana Lewis. Duke and Ripcord are rescued by Scarlett (Rachel Nichols), Snake Eyes (Ray Park), Breaker (Saïd Taghmaoui) and Heavy Duty (Adewale Akinnuoye-Agbaje). They take the warheads to The Pit, G.I. Joe's command center in Egypt, and upon arriving rendezvous with the head of the G.I. Joe Team, General Hawk (Dennis Quaid). Hawk takes command of the warheads and excuses Duke and Ripcord, only to be convinced to let them join his group after Duke reveals that he knows the Baroness. McCullen is revealed to be using the same nanotechnology to build an army of soldiers with the aid of the Doctor (Joseph Gordon-Levitt), planning on using the warheads to cause panic and bring about a new world order. Using a tracking device, McCullen locates the G.I. Joe Base and sends Storm Shadow (Byung-Hun Lee) and the Baroness to retrieve the warheads with assistance from Zartan (Arnold Vosloo). After a fight, Storm Shadow and the Baroness retrieve the warheads and take them to Baron DeCobray, the Baroness's husband, for him to weaponize. Making their way to Paris, the Joes pursue the Baroness and Storm Shadow but are unsuccessful in stopping them from launching the missile. The nanomites destroy the Eiffel Tower and some of the surrounding area before Duke manages to hit the kill switch, but in doing so he is captured and taken to McCullen's base under the Arctic.
The Joes locate the secret base and fly there as McCullen loads three missiles with nano-mite warheads. After Snake Eyes takes out one, Ripcord pursues the remaining missiles in a stolen M.A.R.S. prototype Night Raven jet while Scarlett, Breaker, and Snake Eyes infiltrate the base. Snake Eyes duels and prevails over Storm Shadow. Duke learns that the Doctor is Rex Lewis, Ana's brother believed to have been killed by a mistimed airstrike during a mission led by Duke. Rex had encountered Doctor Mindbender (Kevin O'Connor) in the bunker and became seduced with the nanomite technology, taking too long to retrieve the data and getting caught in the bombing which disfigured him. After freeing Duke, the Baroness gets subdued as the Doctor reveals he has implanted her with nanomites which has put her under his control for the past four years. Attempting to kill Duke, McCullen ends up being burned, so he and the Doctor flee to an escape vessel. Duke and the Baroness pursue him while the Joes fall back when the Doctor activates the base's self destruct sequence.
The Doctor assumes the identity of Cobra Commander, having healed McCullen's burned face with nanomites, encasing it in silver and naming him "Destro", which places McCullen under the Doctor's control. They are captured by G.I. Joe soon after. On the supercarrier USS Flagg, the Baroness is placed in protective custody until they can remove the nanomites from her body. Meanwhile, Zartan, having had his physical appearance altered by nanomites, infiltrates the White House during the missile crisis and assumes the identity of the President of the United States of America (Jonathan Pryce).
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