In the present day, two years after the events of the previous film, Optimus Prime is seen leading NEST, a military organization consisting of human troops and his own team of Autobots (including newcomers Arcee, Chromia, Elita One, Sideswipe, Jolt, and the twins Skids and Mudflap) aimed at killing the remaining Decepticons on Earth. While on a mission in Shanghai, Optimus and his team destroy Decepticons Sideways and Demolishor, being given a warning by the latter that "The Fallen will rise again". Back in the United States, Sam Witwicky finds a splinter of the destroyed AllSpark, and upon contact the splinter fills his mind with Cybertronian symbols. Deeming it dangerous, Sam gives the AllSpark splinter to his girlfriend Mikaela Banes for safe keeping, and leaves her and Bumblebee behind to go off to college. Upon arrival, Sam meets his college roommate Leo Spitz, who runs an alien conspiracy website, and Alice, a co-ed who makes xxxual advances on him. Back home, Decepticon Wheelie tries to steal the shard, only to be captured by Mikaela. After having a mental breakdown, uncontrollably writing in Cybertronian language, Sam calls Mikaela, who immediately leaves to get to him.

Decepticon Soundwave hacks into a US satellite and learns the locations of the dead Decepticon leader Megatron and another piece of the AllSpark. The Decepticons retrieve the shard and use it to resurrect Megatron, who flies into space and is reunited with Starscream and his master, The Fallen in the Nemesis. The Fallen instructs Megatron and Starscream to capture Sam in order to discover the location of the Matrix of Leadership. With Sam's outbreaks worsening, Mikaela arrives at campus just as Alice—revealed to be a Decepticon Pretender—attacks Sam. An irate Mikaela, Sam, and his roommate Leo drive off, destroying Alice, but are seized by the Decepticon Grindor. The Decepticon known as "The Doctor" prepares to remove Sam's brain, but Optimus and Bumblebee turn up and rescue him. In an ensuing fight, Optimus engages Megatron, Grindor and Starscream. Optimus manages to kill Grindor and rip off Starscream's arm, but during a momentary distraction while searching for Sam, he is blindsided then impaled and blasted through the chest by Megatron and dies. Megatron and Starscream depart as the Autobot team arrives to rescue Sam, unable to save Optimus......

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