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Homestuck begins when thirteen-year-old John Egbert receives a beta copy of an upcoming computer game, called "Sburb," in the mail. Installing and running the game on his computer triggers a meteor attack on his real-life house, which survives and is transported to another planet, thus immersing him completely within the world of Sburb. As John's friends Rose, Dave, and Jade join the game with him, they learn that they have unwittingly triggered the end of the world, and that it is their duty to play the game and thereby see the story through to its completion.
As they begin to explore the world of Sburb, John and his friends are harassed by a group of twelve internet trolls who have played the game before. Further contact with the trolls reveals that they are not human at all, but an alien species actually called "trolls". As the trolls gradually become more important to the story, the narrative shifts to a side story exploring the nature of troll society and the specific sequence of events that led to these particular trolls entering the game themselves.
Increased contact between the humans and the trolls leads to an uneasy friendship between the two groups, as well as the discovery that they must cooperate to try to salvage the kids' doomed game session. They learn that the game has a mechanism for restarting, called a "scratch", and they begin working toward the ability to activate it. Fighting against them is a rogue non-player character within Sburb, called Jack Noir, who becomes nearly invincible in their session and is unique because of it, as the trolls had formed an alliance with their version of Jack Noir.
Threatening their plan is a villainous creature called Lord English, a purportedly invincible time-controlling demon, as well as the still-dangerous Jack Noir. The stakes are at their highest, with multiple groups of both humans and trolls forced into a race against time to discover a method of destroying Lord English before he potentially destroys all of existence.