First introduced in July 2002 for high-speed electric passenger services, 53 eight-car units were initially built for use on the West Coast Main Line and leased from Angel Trains. They went to Virgin Trains at first, but in December 2019, they transferred to Avanti West Coast. Four more of the tilting sets were added to the fleet between 2010 and 2012. From 2012, 31 of the original units were increased to 11-coach sets, including 390121, which was decorated in a special ‘Ride Against Climate Change’ livery for Glasgow Cop26 in 2021.