Plot

In May 1943, the crew of the Memphis Belle, a B-17 of the US Army Air Force stationed in England, are grounded while their aircraft is being repaired and they wait for the return of their comrades from a daylight bombing mission over Europe. All of the aircraft arrive back at the airfield, but the last returning aircraft crashes on landing and explodes, killing all aboard. At a party that evening it is announced by Army publicist Lt. Colonel Bruce Derringer (John Lithgow) that the Belle's crew is about to complete its twenty-fifth and final mission. As they will be the first to accomplish this feat in the whole air force, Derringer intends to conduct an elaborate celebration upon their return. His plans are opposed by base commander Colonel Craig Harriman (David Strathairn), who wishes to avoid granting any special treatment to the crew.

The ten-man crew of the Memphis Belle is:

• Captain Dennis Dearborn (Matthew Modine), Pilot. A humorless perfectionist, Dennis worked for his family's furniture business before enlisting. He named the aircraft after his girlfriend. He is openly teased by his crew for his unwavering professional attitude during the before flight checks. he proves to be a competent and capable leader.
• 1st Lt. Luke Sinclair (Tate Donovan), Co-Pilot. The carefree former lifeguard believes himself to be undervalued by Dennis and desires to get his hands on a gun at least once - which he does, with terrible consequences - but proves himself at a critical moment during the mission.
• Lt. Phil Lowenthal (D. B. Sweeney), Navigator. Phil is nervous before their final mission and, convinced that he is doomed to die, drinks too much at the party and distributes his possessions to the rest of the crew.
• Lt. Val Kozlowski (Billy Zane), Bombardier and Medic. While everyone believes the confident, self-assured Val to be a doctor, it is later found that he attended only two weeks of medical school before enlisting.
• Sgt. Danny "Danny Boy" Daly (Eric Stoltz), Radio Operator. An earnest Irish-American, Danny was editor of the school paper, and joined up right after graduation. He is often seen taking pictures of his crew-mates with a small camera which he frequently takes on missions, and writing poetry in a notebook
• Sgt. Virgil "Virgin" Hoogesteger (Reed Diamond), Top Turret Gunner and Flight Engineer. Virge worked for his family's restaurant and plans to open a chain of identical restaurants after the war despite his crew-mate's warnings that such an unheard-of enterprise could never succeed.
• Sgt. Richard "Rascal" Moore (Sean Astin), Ball Turret Gunner. The diminutive Rascal considers himself a ladies' man and delights in teasing his crew mates, especially Virgil. He becomes increasingly nervous of the ball turret's tendency to jam during combat.
• Sgt. Eugene "Genie" McVey (Courtney Gains), Right Waist Gunner. A religious and superstitious Irish-American, Genie carries a St. Anthony's medal on missions with a tendency to lose it. His fellow waist gunner, Jack, often teases him for his superstitious nature, calling him "Genie the Weenie".
• Sgt. Jack Bocci (Neil Giuntoli), Left Waist Gunner. A hot-tempered Chicago hoodlum, Jack appears to look out only for himself but proves surprisingly kind to his fellow waist gunner Genie. He's always eager for a fight, and will often sweet talk to his gun, which he affectionately calls "Mona".
• Sgt. Clay Busby (Harry Connick, Jr.), Tail Gunner. After his father lost the family farm in a poker game, the laconic Clay earned money playing the piano in a New Orleans cat-house. A talented singer, he performs "Danny Boy" at the party before their final mission.

The morning after the party, their target for the day is the city of Bremen, Germany. Experienced officers in the squadron begin to worry. Earlier, it was established that the target was previously attacked a month earlier, and a quarter of the squadron was lost. The Memphis Belle and her squadron are airborne. After assembling their formation and collecting their escort of North American P-51 Mustang fighters, the aircraft fly toward their target only to face constant harassment by defending German Messerschmitt Bf 109 fighters. Eventually the escorting fighters, low on fuel, turn away and the bombers continue alone.

Back at the base, Harriman and Derringer have harsh words as Derringer starts decorating the mess hall for a celebration upon the Belle's return. Derringer accuses Harriman of being concerned only with advancing his personal career. In response, Harriman brings Derringer into his office and angrily dumps a pile of letters on the desk - replies received from families of lost airmen that he himself had written to.

Meanwhile, the bombers have suffered significant losses on their mission. The German interceptors focus their attacks on the aircraft leading the formation. The first lead plane, Windy City loses its engines and explodes right in front of the Belle. More attacks ensue, and the replacement lead plane, the C Cup, is forced to return to base when its nose is crippled by a Bf 109. The crew of the Belle watch in horror when they see a body fall out of the stricken aircraft without a parachute.

Now it is the Belle's turn to take over as lead aircraft of the formation. Finding the target (an aircraft assembly plant) obscured by a smokescreen, Dennis aborts the bomb run and orders the formation to circle again in spite of heavy anti-aircraft fire and the extreme possibility of suffering more casualties to the squadron. Meanwhile, Luke has been itching to do something other than sit in the co-pilot's seat and watch the battle around him. Before they took off he secured an agreement with Clay to fire the tail gunners gun when things got hot. His motives are in the hopes the women back home would find him more desirable if he had actually killed a Nazi. When the moment arrives, Luke get's his wish and shoots down a Messerschmitt. But, his moment of triumph turns to tragedy when his defeated target crashes into the Mother & Country, the same flight of airmen the Belle crew had teased the night before, slicing the bomber in half. In horror and desperation, Danny tries to raise the stricken aircraft on the radio.

The only response heard are their screams of horror, while Luke, struck with immense guilt, watches them plunge to their deaths. On the second run, Val spots the assembly plant through a gap in the smokescreen and the bombers successfully hit their target but, once clear of the anti-aircraft fire, they are again engaged by fighters. Rascal's ball turret jams more than once during the mission, and each time it happens he calls for help from Virgil. The turret is eventually destroyed, but Rascal is spared thanks to his safety strap, and Virgil. The attacks continue, blasting a hole in the fuselage, tearing off a large chunk of the tail, and setting the number four engine on fire. Danny is injured in the attacks, which puts Val in a difficult position when the crew enlists his medical "expertise" to save the injured radioman. Meanwhile, Dennis and Luke skillfully drop the aircraft into a steep dive and put the fire out, despite great risk of losing the aircraft.

Back at base, the remaining squadron aircraft return from their mission, much to the dismay of the ground crews, without the Belle. They wait anxiously for the celebrated aircraft and her crew to return. As the pilots prepare the Belle for landing, Dennis orders them to toss their guns and ammo overboard to lighten their weight, while Val and the enlisted crewmen discuss what to do with Danny. Val, being the only officer with medical knowledge, recommends dumping Danny over German occupied territory believing he'd had have a better chance of survival if found by German soldiers who would take care of him as a prisoner of war, but the remaining crewmen vehemently object doing such an act. Phil interjects and pleads with Val not to throw Danny out, to which he agrees. He cares for Danny all the way back to base.

When Dennis orders the landing gear dropped, only one wheel descends due to electrical failure caused by battle damage. With time running out, Virgil and Jack try to lower the immobile wheel manually with a crank. Soon Phil joins them, having gained new confidence after surviving the mission. He takes over the crank and manages to lower the wheel inches from touchdown. The ground crew, and a slightly humbled Lt. Colonel Derringer, race to greet them with jubilant cheers. The crew exits the battered plane, finally by Dennis, who discovers a bottle of champagne that Virg had smuggled on board. He shows it to the crew, acting as though he is going to reprimand them, and then surprises his crew by joyously cracking the bottle and spraying it in celebration. In the final scene they celebrate their victory with Danny on an ambulance.

The film closes with a dedication to all airmen, friend or foe, who fought in the skies above Europe during World War II.