The story begins with an underachieving fish named Oscar (Will Smith), who is fantasizing about being rich and famous ... while making his way to work as a tongue scrubber at the Whalewash, a business much akin to a car or truck wash. Soon after arriving he is called to the office of his boss, Mr. Sykes (Martin Scorsese), to discuss the fact that he owes "five thousand clams" and has to pay it back by the next day. After explaining this to his best friend Angie (Renée Zellweger) (who harbors a secret love for him), she offers him a chance to pay back the money by pawning off an heirloom pink pearl. Oscar brings the money to the race track to meet Sykes, but becomes distracted by his wishes of grandeur and places it all on a long-shot bet. Such a large bet is noticed nearby by a beautiful fish named Lola (Angelina Jolie), who flagrantly flirts with Oscar until quickly discovering the truth. Sykes is furious that Oscar bet the money but nonetheless agrees to see how the race turns out. Moments before their "horse" crosses the finish line he trips and falls. The race is lost and Oscar is set to be surreptitiously killed.
Meanwhile, on another side of the ocean, a family of criminally-inclined sharks has a problem with one of their sons, Lenny (Jack Black). Lenny refuses to act the part of a killer and wishes to not have to live up to those expectations. With this, his father, Don Lino (Robert De Niro), decides that he must learn to kill, by being taught by Lenny's more savage brother, Frankie (Michael Imperioli).
As the two sharks set out to go in accordance with their father's wishes, Frankie spots the scene where Oscar is being electrocuted by Ernie and Bernie (Doug E. Doug & Ziggy Marley), Sykes' two Jamaican jellyfish thugs, and sends Lenny off to attack. The jellyfish spot Lenny and swim off, leaving Oscar alone with him. Lenny tries to calm Oscar down by telling him that he will not eat him, but Oscar will not listen and continues on screaming. Frankie becomes annoyed and tries to attack, but suddenly an unknown anchor falls and kills him. Lenny is panged with guilt and flees. As no one saw the deed done and Oscar was seen near the body, everyone thinks he did it, and Oscar sees this as the chance to both redeem himself and receive his fame.
Oscar comes back to the city with a new title of the Sharkslayer. Sykes becomes his manager, Lola becomes his girlfriend, and Oscar moves to the "top of the reef" to live in luxury. At the same time Don Lino has everyone out looking for Lenny, and when several get close to Oscar's town the other fish expect him to drive them away. On the way he meets Lenny once more who forces Oscar to let him stay with him because he does not want to go home. Soon Angie finds out about the lie and threatens to tell everyone but Oscar and Lenny convince her to keep quiet.
Although Oscar desires to please everyone, he soon discovers that he pleases no one; his paramour Angie is heartbroken by the fact that Oscar is no longer honest, while her hedonistic rival Lola repeatedly reminds Oscar that he has her only as long as he is famous. With Don Lino planning revenge, Oscar and Lenny stage an event where Lenny pretends to terrorize the town and Oscar must defeat him, throwing him into the depths of the ocean. Though this further cements Oscar as the Sharkslayer, it greatly angers Don Lino. Oscar dumps Lola, to her extreme anger, having never been dumped before. She vows revenge. Oscar buys some Valentine's Day gifts for Angie, but before he can present them to her, he finds that Don Lino has abducted Angie in order to force a meeting. Lenny comes along, now disguised as a dolphin named Sebastian. Angie is bound and gagged and presented to Don Lino on a plate, who prepares to eat her if Oscar doesn't comply. However, Lenny lunges forward and eats Angie up, so Oscar has dominance over the sharks. However, he spends too much time threatening the sharks that he does not realise how much pain Lenny is in, and Angie (along with a lot more scrap metal) is thrown up. Don Lino realizes it's Lenny and pursues Oscar through the reef. Oscar lures both sharks into a trap. Given an ovation by the other fish, Oscar confesses that he is not a Sharkslayer and that it was an anchor that had killed Frankie. However, he lectures Don Lino on. He lectures Don Lino not to prejudge people before he knows them properly and to not make the mistake he made in prejudging his wealth, so Don Lino and Lenny reconcile. Oscar forsakes all the wealth he has acquired, makes peace with the sharks, becomes manager of the Whalewash, and marries Angie. His business grows, as the sharks and whale sharks need to use it as well.
Christina Aguilera and Missy Elliott make a cameo appearance in the end of the movie.
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