Following where the previous film left off, Alice (Milla Jovovich) and the others on the Umbrella Corporation freighter Arcadia face an attack by a fleet of airships led by Alice's former ally, Jill Valentine (Sienna Guillory), who has been brainwashed by Umbrella through a red scarab device attached to her chest. When Alice causes an airship to crash into the ship, the resulting explosion knocks her out and throws her into the water.
The story switches to Alice living as suburban housewife, with her husband Todd (Oded Fehr) and deaf daughter Becky (Aryana Engineer). This idyllic life is disrupted when those undead attack. Alice and Becky escape to the streets and are rescued by a student, Rain Ocampo (Michelle Rodriguez), who lets them ride in her car. As the three escape, they are hit by a truck, knocking Rain unconscious while Alice and Becky escape. Alice runs and is attacked by a plaga infected Todd.
Alice awakens a captive in an Umbrella base. Jill interrogates Alice, who unsuccessfully pleads with her to remember her true identity. During a power failure, Alice escapes from her cell and fights her way through a Tokyo simulation and encounters Ada Wong (Li Bingbing), one of Albert Wesker's (Shawn Roberts) top agents. Ada explains they no longer serve Umbrella. Wesker appears on screen and reveals that the Red Queen is now controlling the entire Umbrella infrastructure and he plans to help Alice escape, in order to save remains of mankind. Ada explains the base is underwater, where they held simulated outbreaks to sell the T-virus to different countries; this led to the world-wide outbreak. To infiltrate the base from outside and help Alice and Ada escape, Wesker has organized a team: Leon S. Kennedy (Johann Urb), Barry Burton (Kevin Durand), and also Luther West (Boris Kodjoe), an Arcadia survivor.
Leon's team plants explosives near the entrance of the base, which will detonate in two hours and trap anybody still inside. The group plans to meet with Alice and Ada in a simulated Raccoon City suburbia, identical to the one in Alice's flashback. In a New York simulation, Alice and Ada defeat two Axemen, prompting the Red Queen to send Jill after them next. Leon and his team enter a Moscow simulation, but are cut off by armed Las Plagas zombies. Alice and Ada manage to enter the suburban simulation where they were supposed to meet up with Leon's team. They find Becky there, revealing the "suburban Alice" is actually a clone of Alice created by Umbrella. Ada explains Umbrella uses the clones for virus outbreak simulations and Umbrella troops; Becky, Todd, and Rain were clones as well. Jill and her mercenaries, consisting of clones of Alice's deceased allies: an "evil" Rain, Carlos Oliviera (Oded Fehr) and James "One" Shade (Colin Salmon) arrive and demand their immediate surrender. A shoot-out occurs, resulting in Ada being captured while Alice and Becky escape.
Alice and Becky run into the "good" Rain and head to Moscow simulation, where they meet up with Leon's team. The group escapes the las plagas, and reach an elevator that leads them to Typhoon-class submarines that could help them escape, although it shuts off. An Uber-Licker appears, with it capturing Becky and killing "good" Rain by breaking her neck. The group pursues the Uber-Licker, where they encounter Jill's group; another battle ensues, in which Barry and One are casualties. Alice manages to rescue Becky despite Leon's discouragement about her just being a clone. During their escape from the Uber-Licker, they arrive at a cloning facility where Alice and Becky see multiple copies of themselves. When Becky asks Alice if she really is her mother, Alice says she is now. The Uber-Licker follows them into the facility, where Alice drops grenades, and fires a grappling hook, escaping the explosion. They rejoin Leon and Luther just as the explosion results in the flooding of the facility.
The group reaches the surface and await pickup but they are met by a submarine, from which Jill, the "evil" Rain, and a captured Ada emerge. With new orders from the Red Queen to kill Alice, Jill attacks her. Rain injects herself with the Las Plagas parasite to give herself superhuman strength and healing; she knocks out Ada and fights Leon and Luther. Alice manages to destroy the scarab device on Jill, returning her to normal; however, unlike Claire, she retains her memories. Rain kills Luther and knocks Leon out. Alice realizes she cannot kill Rain and instead shoots the ice, Rain falls in the water, and vows to get Alice but is dragged under by the Plagas. Leon looks after Ada but Alice collapses from her injuries. She wakes in a helicopter with the other survivors. Alice, Ada, Becky, Leon, and Jill travel to Wesker's base: the heavily barricaded and fortified White House. Wesker injects Alice with the T-Virus, returning her former powers; as she is the only one to successfully bond with it, Alice is humanity's ultimate weapon. On the roof, Wesker explains the Red Queen is trying to destroy all of humanity, and all that remains of the uninfected are in the base; it's humanity's last stand. A pull-away shows Wesker's Umbrella army defending the White House alongside the remaining US Armed Forces against enormous hordes of undead and mutant creatures.
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