Sir Miles Axlerod was voiced by Eddie Izzard who also voices Nigel Bakerbutcher, Professor Zündapp was voiced by Thomas Kretschmann who also portrays Wolfgang von Strucker in the Marvel Cinematic Universe, Grem was voiced by Joe Mantegna who also voices Fat Tony, Acer was voiced by Peter Jacobson, who is best known for portraying Dr. Chris Taub in the FOX medical series, House, Victor Hugo, Vladimir Trunkov, and Ivan were all voiced by Stanley Townsend, J. Curby Gremlin was voiced by John Mainieri, Tubbs Pacer was voiced by Brad Lewis, Alexander Hugo was voiced by Velibor Topic, and Tony Trihull was voiced by Lloyd Sherr.
According to their boss Sir Miles Axlerod, the Lemons have felt rejected by the world because of their defects during the past, an event that left them feel unloved. The film, however, reveals little details regarding how their fellow cars treated them.
As such, Miles Axlerod formulates a plan that involves forcing the entire world to abandon alternative fuel and resorting to using oil (as he still owns the largest oil reserves in the world despite his claims that he has been converted into a hybrid). To that end, Axlerod created the World Grand Prix for racers around the world and a new alternative fuel called Allinol (which is actually gasoline engineered to ignite by an EMP beam). He also plans on murdering anyone who gets in their way.
Being in cahoots with Axlerod's plot and sharing the ownership of the oil reserves, several Lemon families conspired to aid Axelrod in his plan to sabotage the Grand Prix races. To that end, Axelrod gets his right-hand Professor Zündapp to create EMP weapons designed as Prix cameras to ignite any Allinol supply during the races. Because of this, the first two races in Tokyo and Porto Cosa were disasters as many racers engaged into crashes after getting their Allinol ignited by the EMP beams. To cover his tracks, Axlerod suspends on using Allinol for the third and final race in London, though Lightning McQueen stand firm on his decision to use Allinol. Because of this, Axlerod contacts Zündapp to murder McQueen during the final race. This was witnessed by McQueen's best friend Mater, along with two British agents Finn McMissile and Holley Shiftwell, who end up being captured and take hostage in Big Ben.
During the final race, Zündapp gets Grem and Acer to fire up their EMP beam on McQueen, but the latter escapes unharmed (due to Sarge switching McQueen's Allinol supply with Fillmore's organic fuel), just as Mater escapes to warn McQueen of the plot. Anticipating this, Axlerod arranges for Zündapp to place a bomb inside Mater's air filter so that he and McQueen would be killed by the explosion. As Mater runs off out of range, Zündapp ends up being chased by a freed McMissile, forcing the other Lemons to take action and catch McQueen. Though McMissile and Holley manage to defeat Zündapp, the other Lemons surround them in an intersection, preparing to wait for the imminent explosion to kill McQueen, Mater and the agents. Mater tries to talk the Lemons out of the plan by pointing out that it won't make them feel any better, but even though the Lemons agree with him, they still want him and his friends dead for their greed. Fortunately, McQueen brought in his friends from Radiator Springs to stop them, forcing the Lemons to surrender.
Mater eventually proceeded to expose Axlerod as the mastermind behind the plot by forcing him to deactivate the bomb within close range. With the plot exposed, the Lemons are arrested for conspiracy to commit crime, murder, attempted murder, and embezzlement.
- Sir Miles Axlerod (leader, arrested)
- Professor Zündapp (second-in-command, arrested)
- Grem and Acer (lieutenants, fate unknown)
- J. Curby Gremlin (leader; arrested)
- Tyler Gremlin (fate unknown)
- Don Crumlin (fate unknown)
- Towga Gremlin (deceased)
- Stefan Gremsky (fate unknown)
- Tubbs Pacer (leader; arrested)
- Petey Pacer (arrested)
- Fred Pacer (deceased)
- Victor Hugo (leader; arrested)
- Alexander Hugo (fate unknown)
- Ivan (arrested)
- Karl Haulzemoff (fate unknown)
- Vladmir Trunkov (leader; arrested)
- Petrov Trunkov (fate unknown)
- Tolga Trunkov (arrested)
- Tony Trihull (deceased)
- Jerome Ramped (fate unknown)
- Muggsy Liftsome (fate unknown)
- Though they are considered Cars' darker villains due to their willingness to commit murder, Mater sympathizes with them due to their troubled pasts. That said, the narrative and Mater himself make it clear that it doesn't justify their crimes. Furthermore, even some of their more ruthless members agree that it wouldn't make them feel better, implying that they are killing people out of both greed and resentment.