Thousands of years before the events of the first film, the Dynasty of Primes traveled through the galaxy to create Energon by creating sun-absorbing machines called Sun Harvesters. The Dynasty of Primes followed a law that no sun may be destroyed where there was life on one of its planets. However, one of the Primes felt hatred towards the human race and attempted to break this law by building a Sun Harvester on Earth in 17,000 B.C . The rest of the Dynasty named himThe Fallen, and sacrificed themselves to hide the Sun Harvester's key, the "Matrix of Leadership".
Two years after the first film, Sam Witwicky is heading off to college. While packing, he finds a shard of the All Spark in the clothing he was wearing during the fight. Unbeknownst to Sam, when he touches the shard, it "uploads" its knowledge into his mind. Sam gives the shard to his girlfriend Mikaela Banes for safekeeping, and leaves his guardian Bumblebee behind.
The Autobots and the humans have created NEST (Network Elements: Supporters and Transformers), an elite joint military organization from multiple countries from the top elite soldiers on the planet led by Optimus Prime, Major William Lennox and General Morshower. In Shanghai, Optimus takes down Demolishor who warns him of The Fallen's return. Reporting to NEST's base in Diego Garcia, the Autobots are patronized by U.S. National Security Advisor Galloway for their actions. Soundwave, a Decepticon communications officer, hacks into a U.S military satellite, and overhears both the location of Megatron's corpse (buried at sea in the Laurentian Abyss) and the presence of a shard of the Allspark stored at the base. Soundwave deploys Ravage to retrieve the shard. The Constructicons, Ravage and the mad surgeon known as "The Doctor" dive into the Laurentian Abyss and resurrect Megatron with the salvaged shard. Megatron flies to the Decepticon flagship Nemesis, where The Fallen rests and Starscream has awaited Megatron's arrival. The Fallen instructs Megatron to kill Optimus and capture Sam, as he has knowledge that can lead to the Matrix of Leadership.
At her shop, Wheelie tries to steal Mikaela's shard, but she captures him in the process and heads to see Sam with Wheelie locked in a container. At college, Sam meets his roommate Leo Spitz, who runs a conspiracy website. During class, Sam experiences a flash of Cybertronian symbols in his mind. He has a meltdown in class and starts graffiting the symbols all around his room. Mikaela arrives just as Sam is attacked by a Pretender. Sam, Mikaela and Leo escape, but are quickly captured by the Decepticons torturing Sam to obtain the information he has about the whereabouts of the Matrix. Optimus and Bumblebee arrive and battle Megatron, Starscream, and Grindor. Grindor is killed, Starscream is wounded and Megatron then kills Optimus by impaling Optimus's chest and firing. Bumblebee and the twins, Skids and Mudflap, flee with the humans. The NEST team extracts the body of Optimus and returns it to New Jersey base until a convoy of the U.S. Army arrives with weapons all aimed at the Autobots. Lennox tells them to stand down, but Galloway arrives with the news that NEST has been decommissioned.
Meanwhile, The Fallen taps into media signals and broadcasts a demand that the people of Earth deliver Sam to the Decepticons. The Decepticons then also kidnap Sam's parents. Leo suggests to Sam and Mikaela that his online rival "RoboWarrior" may be able to help. When the trio find him, they discover that RoboWarrior is Agent Simmons from the now defunct Sector Seven (as seen in the 2007 Transformers film). Simmons reveals Transformers have been on Earth for a millennium, and convinces Wheelie to reveal a group of ancient Transformers who live on Earth, known as the Seekers. They find the elderly Jetfire who teleports them to Egypt and reveals the history of the Transformers.
Following several clues, the trio find the Tomb of the Primes in Petra. Led to an ancient Egyptian temple, the group discovers the Tomb, with the Matrix however crumbling to dust upon Sam touching it. Sam puts the sand in a pouch to take with him. Simmons contacts Lennox who flies over with his troops and the Autobots. The Decepticons arrive and battle the Autobots and NEST, who are aided by Egyptian and Jordanian military. Constructicon Devastator uncovers the Sun Harvester from inside a pyramid before being killed by a rail gun strike ordered by Simmons. The Air Force is called in for an airstrike from every aircraft in their fleet, wiping out the Decepticons, leaving only Megatron and Starscream. Sam makes a break for it towards Optimus until he is killed by Megatron. In a limbo between life and death, Sam has a vision of the Dynasty of the Primes who tell him that the Matrix of Leadership is not a physical item, but a right that he has earned.
Sam awakens, and the Matrix reforms from the sand in Sam's pouch. Sam uses the Matrix to resurrect Optimus Prime, but The Fallen arrives and retrieves the Matrix in order to activate the Sun Harvester. The U.S. military forces attack the Sun Harvester, but are quickly wiped out by The Fallen's telekinesis. Combined with Jetfire who willingly sacrifices his Spark and parts, Optimus destroys the Sun Harvester, kills The Fallen, and wounds Megatron. Starscream convinces Megatron to retreat. Sam and Mikaela profess their love for each other, and Optimus thanks Sam for his bravery and records another message in his data bank saying that the humans and the Transformers share a common past. During the credits, Sam is seen returning to college.
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