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Two weeks before her maiden voyage, Charles boarded the Titanic and acted as First Officer for the sea trials. Captain Edward J. Smith made Henry T. Wilde, of the Olympic, the Titanic's Chief Officer causing the original Chief Officer William McMaster Murdoch to drop to First Officer and Lightoller to Second Officer. The original Second Officer David Blair was dropped from the voyage altogether while the ship's roster of junior officers remained unchanged.

On the night of 14 April 1912, Lightoller commanded the last bridge watch prior to the ship's collision with an iceberg before being relieved by Murdoch. Lightoller had retired to his cabin and was preparing for bed when he felt the collision occur. Wearing only his pajamas, Lightoller hurried out on deck to see what had happened but after seeing nothing retired back to his cabin. Figuring it would be better to remain where other officers knew where to find him if they needed him, he lay awake in his bunk until Fourth Officer Boxhall summoned him to the bridge. He pulled on trousers and a Navy-blue sweater over his pajamas and also donned (along with socks and shoes) his officers' overcoat and hat.

Once the fate of the ship became clear, Lightoller immediately went to work assisting in the evacuation of the passengers into the lifeboats. Lightoller was notably stricter than some of the other officers in observing the rule of "women and children first", almost to the point of the rule being "women and children only." Lightoller lowered the lifeboats on the port side of the Titanic. The officer's last action was attempting to launch Collapsible B which was a smaller, Englehardt lifeboat with canvas sides that was stowed atop the officers' quarters on the hurricane deck on the port side. As the ship sank, sea water washed over the entire bow of the Titanic; producing a large wave that rolled aft along the boat deck. Seeing crowds of people run away from the rising water and the collapsible boat wash away upside down, Lightoller decided not to prolong it and dove into the water.

Once surfaced from his dive, he spotted the ship's crow's nest now level with the water and temporarily swam towards it as a place of safety before realizing that is was safer to clear away from the foundering vessel. Then Lightoller was sucked under as water flooded down one of the forward ventilaters. He was pinned there against the grating for a few seconds. Luckily, a blast of hot air from the depths of the ship erupted out of the ventilater and blew him to the surface. Following this, the officer saw Collapsible B, which the crew had unsuccesfully tried to launch earlier, floating upside down with several swimmers clawing to it. He stroked to it and held on by a rope at the front.

Then one of the Titanic's massive funnels broke free and hit the water, washing the collapsible further away from the sinking ship. Later on, Officer Lightoller took charge and was able to calm and organize the survivors (numbering around thirty) who were on the overturned lifeboat. He led them in yelling in unison "Boat ahoy!" but with no success. During the night the sea began to rise and Lightoller led the men in shifting their weight with the swells so that their craft would not be swamped. Had they not done this, they would have been thrown into the frigid water again. The men kept this up at his direction for hours in the freezing weather until they were finally rescued by another lifeboat. Second Officer Lightoller was the last survivor to come aboard the rescue ship Carpathia.

As the senior surviving crew member, Lightoller was a key witness at both the American and British inquiries. He blamed the accident on the sea that night being the calmest he ever saw in his life; the floating icebergs gave no tell tale early warning signs of breaking whitewater at their base. He deftly defended his employer the White Star Line despite hints of excessive speed, missing binoculars in the crows' nest, and the plain recklessness of traveling through an ice field on a calm night when all other ships in the vicinity thought it wiser to stop until morning. Lightoller was also able to help channel public outcry over the incident into positive change as many of his recommendations for avoiding such accidents in the future were adopted by maritime nations. Basing lifeboat capacity on numbers of passengers and crew instead of ship tonnage, lifeboat drills so passengers know where their lifeboats are and crew know how to operate them, manned 24 hour wireless (radio) communications in all passenger ships, and official ice warnings from the maritime board are some of his recommendation made at the inquires.