His family released a statement Sunday morning to announce his death: "His loss leaves a void in our lives that will never be filled."
"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."
Paterno's son Scott said on Nov. 18 that his father was being treated for lung cancer. The cancer was diagnosed during a follow-up visit for a bronchial illness. A few weeks after that revelation, Paterno also broke his pelvis after a fall but did not need surgery.
Paterno had been in the hospital since Jan. 13 for observation for what his family had called minor complications from his cancer treatments. Not long before that, he conducted his only interview since losing his job, with The Washington Post. Paterno was described as frail then, speaking mostly in a whisper and wearing a wig. The second half of the two-day interview was conducted at his bedside.
"As the last 61 years have shown, Joe made an incredible impact," said the statement from the family. "That impact has been felt and appreciated by our family in the form of thousands of letters and well wishes along with countless acts of kindness from people whose lives he touched. It is evident also in the thousands of successful student athletes who have gone on to multiply that impact as they spread out across the country."
The final days of Paterno's Penn State career were easily the toughest in his 61 years with the university and 46 seasons as head football coach.
It was because Paterno was a such a sainted figure -- more memorable than any of his players and one of the best-known coaches in all of sports -- that his downfall was so startling. During one breathtaking week in early November, Paterno was engulfed by a scandal and forced from his job, because he failed to go to the police in 2002 when told a young boy was molested inside the football complex.
"I didn't know which way to go ... and rather than get in there and make a mistake," he said in the Post interview.
Jerry Sandusky, the former defensive coordinator expected to succeed Paterno before retiring in 1999, was charged with sexually assaulting 10 boys over 15 years. Two university officials stepped down after they were charged with perjury following a grand jury investigation of Sandusky. But attention quickly focused on an alleged rape that took place in a shower in the football building, witnessed by Mike McQueary, a graduate assistant at the time.
McQueary testified that he had seen Sandusky attacking the child and that he had told Paterno, who waited a day before alerting school authorities. Police were never called and the state's top cop later said Paterno failed to execute his moral responsibility by not contacting police.
"You know, (McQueary) didn't want to get specific," Paterno said in the Post interview. "And to be frank with you I don't know that it would have done any good, because I never heard of, of, rape and a man. So I just did what I thought was best. I talked to people that I thought would be, if there was a problem, that would be following up on it."
On the morning of Nov. 9, Paterno said he would retire following the 2011 season. He also said he was "absolutely devastated" by the abuse case.
"This is a tragedy," the coach said. "It is one of the great sorrows of my life. With the benefit of hindsight, I wish I had done more."
But the university trustees faced a crisis, and in an emergency meeting that night, they fired Paterno, effective immediately. Graham Spanier, one of the longest-serving university presidents in the nation, also was dismissed.
According to Lanny Davis, an attorney retained by the trustees as an adviser, board vice chairman John Surma regretted having to tell Paterno the decision over the phone.
The university handed the football team to one of Paterno's assistants, Tom Bradley, who said Paterno "will go down in history as one of the greatest men, who maybe most of you know as a great football coach."
Thick, smoky-lens glasses, rolled up khakis, jet-black sneakers, blue windbreaker -- Paterno was easy to spot on the sidelines. His teams were just as easy to spot on the field; their white helmets and classic blue and white uniforms had the same old-school look as the coach.
Paterno believed success was not measured entirely on the field. From his idealistic early days, he had implemented what he called a "grand experiment" -- to graduate more players while maintaining success on the field.
He was a frequent speaker on ethics in sports, a conscience for a world often infiltrated by scandal and shady characters.
His teams consistently ranked among the best in the Big Ten for graduating players. As of 2011, it had 49 academic All-Americans, the third-highest among schools in the Football Bowl Subdivision. All but two played under Paterno.
"He teaches us about really just growing up and being a man," former linebacker Paul Posluszny, now with the NFL's Jacksonville Jaguars, once said. "Besides the football, he's preparing us to be good men in life."
Paterno certainly had detractors, as well. One former Penn State professor called his high-minded words on academics a farce. He was criticized for making broad critiques about the wrongs in college football without providing specifics. A former administrator said his players often got special treatment compared to non-athletes. His coaching style often was considered too conservative. Some thought he held on to his job too long. There was a push to move him out in 2004 but it failed.
But the critics were in the minority, and his program was never cited for major NCAA violations. However, the child sexual abuse scandal prompted separate investigations by the U.S. Department of Education and the NCAA into the school's handling.
Paterno played quarterback and cornerback for Brown University and set a defensive record with 14 career interceptions, a distinction he boasted about to his teams all the way into his 80s. He graduated in 1950 with plans to go to law school. He said his father hoped he would someday be president.
When he was 23, a former coach at Brown was moving to Penn State to become the head coach and persuaded Paterno to come with him as an assistant.
"I had no intention to coach when I got out of Brown," Paterno said in 2007 at Beaver Stadium in an interview before being inducted into the Hall of Fame. "Come to this hick town? From Brooklyn?"
In 1963, he was offered a job by the late Al Davis -- $18,000, triple his salary at Penn State, plus a car to become general manager and coach of the AFL's Oakland Raiders. He said no. Rip Engle retired as Penn State head coach three years later, and Paterno took over.
At the time, the Lions were considered "Eastern football" -- inferior -- and Paterno courted newspaper coverage to raise the team's profile. In 1967, PSU began a 30-0-1 streak.
But Penn State couldn't get to the top of the polls. The Lions finished second in 1968 and 1969 despite perfect records. They went 12-0 in 1973 and finished fifth. Texas edged them in 1969 after President Richard Nixon, impressed with the Longhorns' bowl performance, declared them No. 1.
"I'd like to know," Paterno said later, "how could the president know so little about Watergate in 1973, and so much about college football in 1969?"
A national title finally came in 1982, in a 27-23 win over Georgia at the Sugar Bowl. Penn State won another in 1986 after the Lions picked off Vinny Testaverde five times and beat Miami 14-10 in the Fiesta Bowl.
They have made several title runs since then, including a 2005 run to the Orange Bowl and an 11-1 campaign in 2008 that earned them a berth in the Rose Bowl, where they lost 37-23 to Southern California.
In his later years, physical ailments wore the old coach down. Paterno was run over on the sideline during a game at Wisconsin in November 2006 and underwent knee surgery. He hurt his hip in 2008 demonstrating an onside kick.
An intestinal illness and a bad reaction to antibiotics prescribed for dental work slowed him for most of the 2010 season. Paterno began scaling back his speaking engagements that year, ending his summer caravan of speeches to alumni across the state.
Then a receiver bowled over Paterno at practice in August, sending him to the hospital with shoulder and pelvis injuries and consigning him to coach much of the season from the press box.
"The fact that we've won a lot of games is that the good Lord kept me healthy, not because I'm better than anybody else," Paterno said two days before he won his 409th game and passed Eddie Robinson of Grambling State for the most in Division I. "It's because I've been around a lot longer than anybody else."
Paterno could be conservative on the field, especially in big games, relying on the tried-and-true formula of defense, the running game and field position.
"They've been playing great defense for 45 years," Iowa coach Kirk Ferentz said in November.
Paterno and his wife, Sue, raised five children in State College. Anybody could telephone him at his modest ranch home -- the same one he appeared in front of on the night he was fired -- by looking up "Paterno, Joseph V." in the phone book.
He walked to home games and was greeted and wished good luck by fans on the street. Former players paraded through his living room for the chance to say hello. But for the most part, he stayed out of the spotlight.
Paterno did have a knack for joke. He referred to Twitter, the social media, as "Twittle-do, Twittle-dee."
He also could be abrasive and stubborn, and had his share of run-ins with his bosses or administrators. And as his legend grew, so did the attention to his on-field decisions, and the questions about when he would retire.
Calls for his retirement reached a crescendo in 2004. The next year, Penn State went 11-1 and won the Big Ten. In the Orange Bowl, PSU beat Florida State, whose coach, Bobby Bowden, left the Seminoles after the 2009 season after 34 years and 389 wins.
Like many others, he was outlasted by "JoePa."