His family released a statement Sunday morning to announce his death.
"He died as he lived," the statement said. "He fought hard until the end, stayed positive, thought only of others and constantly reminded everyone of how blessed his life had been. His ambitions were far reaching, but he never believed he had to leave this Happy Valley to achieve them. He was a man devoted to his family, his university, his players and his community."
The winningest coach in Division 1 football was dismissed from his post in November during the aftermath of the child sex abuse charges against former assistant coach Jerry Sandusky.
Shortly after his ouster, Paterno was diagnosed with lung cancer and broke his pelvis, the same injury he sustained during preseason practice last year.
Paterno served as head coach for the Penn State Nittany Lions football team for 46 seasons. He won 409 games and took his team to 37 bowl games and two national championships. More than 250 of the players he coached went on to the NFL.
He leaves behind his wife, five children and 17 grandchildren.
In a statement released Sunday, Sandusky called the death of his former boss a sad loss.
"This is a sad day! Our family, Dottie and I would like to convey our deepest sympathy to Sue and her family," Sandusky said in the statement. "Nobody will be able to take away the memories we all shared of a great man, his family, and all the wonderful people who were a part of his life."
Sandusky is awaiting trial on 52 criminal counts for what prosecutors say was sexual abuse of 10 children over 15 years. He denies the allegations.
Sandusky was Paterno's top assistant for years until he retired in 1999, and he said he remembered Paterno as a great man who met high standards in a difficult job.
"He maintained a high standard in a very difficult profession. Joe preached toughness, hard work and clean competition," Sandusky said. "Most importantly, he had the courage to practice what he preached."
See photos of former Penn State head football coach Joe Paterno through the years.
ABC News and The Associated Press contributed to this report.
