He was a frequent speaker on ethics in sports, a conscience for a world often infiltrated by scandal and shady characters.
The team 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. One former Penn State professor called his high-minded words on academics a farce, and a former administrator said players often got special treatment. His coaching style often was considered too conservative. Some thought he held on to his job too long, and a move to push him out in 2004 failed.
But the critics were in the minority, and his program was never cited for major NCAA violations. The child sex abuse scandal, however, did prompt separate investigations by the U.S. Department of Education and the NCAA into the school's handling.
Paterno played quarterback and defensive back for Brown University and set a defensive record with 14 career interceptions, a distinction he still boasted about to his teams in his 80s. He graduated in 1950 with plans to go to law school. He said his father hoped he would someday be president.
But when Paterno 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 in an interview at Penn State's Beaver Stadium before being inducted into college football's 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 seasons. They were undefeated and untied again in 1973 at 12-0 again but 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, after a 27-23 win over Georgia at the Sugar Bowl. Another followed in 1986 after the Lions picked off Vinny Testaverde five times and beat Miami 14-10 in the Fiesta Bowl.
They made several title runs after that, including a 2005 run to the Orange Bowl and an 11-1 season in 2008 that ended in a 37-23 loss to Southern California in the Rose Bowl.
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. He 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 what would be his last 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 jokes. He referred to Twitter, the social media site, as "Twittle-do, Twittle-dee."
He also could be abrasive and stubborn, and he 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 hang it up.
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, was eased out after the 2009 season after 34 years and 389 wins.
Like many others, he was outlasted by "JoePa."