Sixty-five million years ago, an alien race known as the "Creators" used devices called Seeds to terraform planet Earth, effectively wiping out the dinosaurs and covering it with an alloy called Transformium. In the present day, geologist Darcy Tirrel excavates the Transformium for K.S.I. Industries, which uses it to build man-made artificial Transformer drones.
Five years after the Battle of Chicago, the humans have begun to view the Transformers as a threat to public safety, leading the U.S government to terminate all human-Autobot joint programs. Although the public believes that the Autobots have been granted sanctuary, they are actually being hunted by a CIA black ops division known as Cemetery Wind led by opportunistic government official Harold Attinger, who believes that all Transformers should be exterminated regardless of faction. They are aided by Lockdown, a Cybertronian bounty hunter working for the Creators, who promises to give Attinger a Seed if his division manages to capture Optimus Prime. Cemetery Wind locates Ratchet in Mexico City and Lockdown kills him when he refuses to give up Optimus' whereabouts.
Optimus, damaged and disguised as a rundown semi-truck, is discovered in an abandoned theater by Cade Yeager, a financially struggling Texan inventor, and brings him back to his farm. While his teenage daughter Tessa and business partner Lucas Flannery encourage him to turn Optimus over to the authorities after realizing what he really is, Cade instead fixes Optimus, hoping to understand his technology, and unexpectedly revives him. Still skeptical of Optimus, Lucas calls Cemetery Wind, which attacks and destroys the farm, but Optimus and Tessa's boyfriend, Irish rally car driver Shane Dyson, come to their aid. While on the run from Cemetery Wind and Lockdown, Lucas is killed by one of Lockdown's grenades. They flee to the desert and take refuge in an abandoned gas station. Later, Optimus scans a new, more functioning alternate mode, then summons the surviving Autobots – Bumblebee (who now has an alternate form), Hound, Drift, and Crosshairs. Using a CIA drone that he stole during the attack, Cade discovers K.S.I.'s involvement in the attacks on the Autobots. Optimus vows to personally kill Attinger for his actions against his brethren.
Infiltrating K.S.I.'s headquarters in Chicago, Cade discovers the remains of Autobots and Decepticons are being melted down to make the drones. Joshua Joyce, the ambitious company CEO, is collaborating with Attinger to revolutionize global defenses and improve human society using the Seed. He has captured the Autobot Brains and is using him and Megatron's head to create prototype Transformer soldiers named Galvatron and Stinger. Outraged at Cade's discovery, the Autobots storm the building (in which Bumblebee gains a different alternate form along the way), rescue Brains, and destroy the laboratory, but they leave after Joshua declares they are no longer needed in the world. Attinger forces Joshua to deploy Galvatron and Stinger to attack the Autobots. During the battle, Galvatron starts destroying vehicles on its own will and even speaks to Optimus while fighting him. Lockdown then arrives and abducts Optimus while Galvatron and Stinger retreat; Tessa is inadvertently captured as well.
While Lockdown's large prison spacecraft hovers over Chicago to hand over the Seed, Cade, Shane, and the Autobots use this opportunity to sneak on board and rescue Optimus and Tessa. They hijack a smaller ship, containing a number of other Autobots called the Dinobots, just before Lockdown leaves Earth. The Autobots reveal to Cade that Galvatron is, in fact, Megatron, who transferred his consciousness into the Galvatron drone after K.S.I. officials accidentally revived him. They realize that Megatron is plotting to use the Seed and the K.S.I. drones to conquer the world, by detonating the Seed in Hong Kong, as K.S.I. has brought it there to use it in the remote Mongolian desert to create vast amounts of usable Transformium. Cade informs Joshua of Megatron's presence, causing him to have a change of heart and agreeing to hand over the Seed with help from Darcy and his Chinese business associate Su Yueming. Meanwhile, Lockdown finds out about Optimus's escape and returns to Earth to retrieve him and the Dinobots.
Optimus decides the Autobots should no longer help humans since they revived Megatron, but Cade convinces Optimus to not lose faith in them. Back at the facility, Megatron reactivates himself and takes control of the K.S.I. drones. A battle breaks out in Hong Kong's streets between Cade's group, the Autobots, Cemetery Wind, Megatron, and his drones. During the fight, Optimus forms an alliance with the Dinobots, who help them fight the drones while Bumblebee kills Stinger. Lockdown returns to recapture Optimus and the Dinobots, using a large magnet in his ship that picks up and drops metal objects, causing destruction to the city. After disabling the magnet, Optimus fights Lockdown at an abandoned factory. In the ensuing duel, Optimus kills Attinger (who still considers all Cybertronians as a threat) to save Cade before being impaled with his own sword by Lockdown. Cade and Bumblebee fight Lockdown while Tessa and Shane use a tow truck to free Optimus, who kills Lockdown before using one of his grenades to finish off the remaining drones. Megatron retreats, vowing to return.
As Lockdown's ship leaves Earth, Optimus tells the Autobots to protect the Yeagers and lets the Dinobots go. He then flies away into space with the Seed to hide it away, while sending a message to the Creators that he is coming for them.
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