In the beginning of the game, Crash Bandicoot aids Coco with a butter-recycling device.[10] This peace is interrupted by Doctor Neo Cortex, who arrives in his airship. Cortex captures Coco and Aku Aku, and encases Crunch in crystal. In an attempt to stop Doctor Cortex, Crash throws Coco's machine at the airship, and manages to sever Aku Aku's cage, which falls into the nearby forest. When Crash rescues Aku Aku, the latter explains that his powers are useless against the fierce mutants that they have encountered, and suspects that dark magic is involved.[11] When Crash and Aku Aku follow Cortex's blimp to see what he's up to, they discover that Cortex and Uka Uka are stealing all the mojo from the Temple of Zoom, an event somehow linked to the strange mutants that have appeared.[12] Upon reaching the roof of the Temple, Neo Cortex reveals his plot to use the stolen Mojo to create a huge army of loyal mutants, and use those mutants to build a colossal robot known as the Doominator, which will crush the Wumpa Islands and take over the world.[13] Uka Uka then leaves Cortex and his patented Yuktopus to deal with Crash and Aku Aku while he takes the Mojo and Coco back to their base.[14] After the Yuktopus fails to defeat Crash, Cortex boasts that Crash will never find their base, and flies off, leaving Crash and Aku Aku to follow him.[15]
Back at the Cortex base, Uka Uka derides Cortex for failing to destroy Crash, dismissing his antics as "ridiculous and appalling", and decides to replace him with his niece, Nina Cortex, despite protests from Neo, N. Gin, and Tiny Tiger.[16] Nina's first act as the main antagonist is to have Coco brainwashed and have her aid in the construction of the Doominator.[17] Meanwhile, Crash and Aku Aku come across a giant wood-cutting/mining area, where logging machinery and giant digging robots reign supreme, and all the trees in the would-be jungle are gone. After working their way through the logging areas, Crash and Aku Aku go into a tunnel that heads into a lake of lava surrounded by mountains. On the other side are two giant metallic mosquitoes, which are using drills to suck the minerals from the lava inside the volcanic area. After the two mosquitoes are destroyed, Tiny Tiger, who is behind the mining operations, literally jumps in to express his frustration with Crash for hindering his job and for not inviting him to Crash Tag Team Racing.[9][18] After a quick showdown of sorts, Crash uses a Shellephant to force Tiny to tell them where Coco has been taken. Tiny spills the beans that she is being held at the factory on the beach, and that Nina has taken over the leadership role, to which Aku Aku and Crash display much disbelief. Upon realizing that this is the truth, Aku Aku and Crash head to the beach in a hurry.[19]
Crash and Aku Aku arrive at the beach and begin to approach the factory. Crash, who realizes just how far away the factory is, throws a large temper tantrum, eventually moving on, having accepted that he must save his sister. After a long trek through the beach and its surrounding jungle area, Crash and Aku Aku move through a sewer line, finally arriving outside the factory, which appears to be modelled after N. Gin. Finally infiltrating the factory, Crash confronts N. Gin in the head of the Statue of Liberty-like base. After a quick showdown, Aku Aku demands information from N. Gin, who responds by having a conversation between two of his conflicting beliefs: one part of N. Gin wants Neo Cortex to be freed, while the other is happy with Nina's new way of ruling. Eventually, they reach a compromise in which N. Gin will tell Crash and Aku Aku where the others are, believing they will help Cortex.[20] Aku Aku once more demands information, and N. Gin delivers, stating that the area they seek is in Uka Uka's lab, which is located on the giant tree in the center of Wumpa Islands.[21]
Aku Aku, knowing the island best, warps himself and Crash to the tree, which is more infested with Titans than any other area. Uka Uka senses Aku Aku's presence through the Mojo, and stays in the lab while Nina goes to work on the Doominator.[22] Arriving in the middle of the tree, Aku Aku notices a giant house, with flashing lights coming out of all the windows, and electricity spewing from the bottom. The duo enters the house, where they encounter Uka Uka in the furthest most room. Uka Uka floats towards the top of the room, where he points out the large electric beams flowing towards a familiar device. Uka Uka asks Crash if he remembers the device, noting its name as the Evolvo-Ray, used by Doctor Neo Cortex to create Crash in the first game of the series. Uka Uka then fires the ray upon himself, which causes a body of wood and vines to grow from the back of his mask.[23] Uka Uka faces his defeat soon after, and claims he'll have the last laugh, as Nina Cortex is already about to launch the Doominator robot. Aku Aku disregards this, telling Crash that he senses Coco, and can warp them to the Doominator.[24]
Upon doing this, the duo arrives at the bottom of the Doominator. They enter the base, which is similar to a giant museum-like mansion. At the end of the mansion is an entrance to the giant dome that rests on the Robot's head. The dome is filled with molten metal produced from the processes that run the Doominator. After Crash and Aku Aku pass this area, they finally arrive at the face of the Robot, which resembles Neo Cortex. The two enter the robot, and are confronted by Nina Cortex. Nina finally releases her uncle, commanding him to keep the robot on course with Wumpa Island itself, which the Cortexes plan to destroy. Cortex responds by ordering Crash to "take out this terrible excuse for a niece!"[25] Nina summons her Arachnina robot, and fights Crash.[26] Crash eventually overcomes the robot, and destroys it. Coco is freed, and the Doominator can be disabled. The collapsing Doominator just barely misses the Bandicoot home (and Crunch), sparing much of Wumpa Island. Escaping the collapsed Doominator, Cortex praises Nina for betraying him, and promises to be more evil in the near future, but proclaims that he's still going to spank Nina stupid for all of this. The Bandicoots decide that it is time for celebration, leading Crash to shout his first word and the object of their celebration: pancakes.[27]
The Nintendo DS version of the game largely follows the same story of the home console versions, but with minor tweaks involved, such as Nina Cortex having a more predominant role and the inclusion of Dingodile.[6] The Game Boy Advance version also includes Dingodile, and has some plot differences, such as the Arachnina (reffered to by its old name, "Spiderbot") not being the last boss, and Cortex being reinstated after Nina's defeat in the N. Trapment Island (the penultimate island in the game).
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