Gurren Lagann takes place in a fictional future in which human beings have been forced under Earth's surface and live in isolated subterranean civilizations. These "villages" have no contact with the surface world or other villages. Because frequent earthquakes damage infrastructure, the villages must constantly extend deeper into the earth — individuals relegated to this task are known as "diggers".

Jeeha Village is the home of Simon, a fourteen-year-old digger who was orphaned when his parents were killed in an earthquake. Although Simon is respected by the village elders as the best of the diggers, he is ostracized by his peers. Simon discovers a special drill key deep in the ground and is soon recruited by an eccentric fellow orphan named Kamina into his group "The Gurren Brigade". Kamina dreams of the surface world, which he once visited as a child, though he had to return home as he was too weak to survive there.

After a failed attempt at reaching the surface, Kamina is jailed by the village elder. Simon resumes digging, only to discover "a big face" deep in his tunnel. As he excitedly fetches Kamina to show him the discovery, Simon is also caught by the village elder. Fortuitously, Kamina's punishment is interrupted by the collapse of the ceiling over Jeeha Village, as an enormous robot falls into the cavern. A girl named Yoko appears and tries to defeat the machine with a large rifle, but to no avail. Simon takes Kamina and Yoko to the "big face" he found earlier and discovers that the small drill key can be used to activate a small robot, named "Lagann", which he uses to destroy the invading robot and effectively break through to the surface.

Once on the surface, Simon and Kamina discover there are more large robots known as "Gunmen" (piloted by "Beastmen") that terrorize the surface dwellers during the day. Simon and Kamina help Yoko and her village destroy three more Gunmen of which Kamina takes one for himself, naming it the "Gurren". Eventually, Simon and Kamina decide to embark on a journey to end the constant struggle between the Beastmen and humans, and are initially accompanied by Yoko and Leeron, a mechanic from Yoko's village. As they head out to find the headquarters of the Beastmen's Human Eradication Army, the group encounters various Beastmen warriors including the vicious Viral.

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