State College, Pennsylvania (CNN) -- Joe Paterno, whose tenure as the most successful coach in major college football history ended abruptly in November amid allegations that he failed to respond forcefully enough to a sex abuse scandal involving a former assistant, died Sunday, a family spokesman said. He was 85.
The longtime Penn State head coach was diagnosed with what his family had called a treatable form of lung cancer shortly after the university's Board of Trustees voted to fire him.
He had been hospitalized in December after breaking his pelvis in a fall at his home and again in January for what his son called minor complications from his cancer treatments.
"It is with great sadness that we announce that Joe Paterno passed away earlier today," the family statement said. "His loss leaves a void in our lives that will never be filled."
Paterno, who was affectionately known as "JoePa" by generations of his players and football fans alike, was widely admired in football circles for what he called his "Grand Experiment" -- his expectation that big-time college football players could succeed on the field while upholding high academic and moral standards away from the gridiron.
Under his leadership, the Nittany Lions won two national championships, went undefeated five times and finished in the top 25 national rankings 35 times, according to his official Penn State biography.
At the same time, the program never fell under NCAA sanctions for major infractions while producing 13 Academic All-Americans since 2006. In 2009, according to the university, the Nittany Lions posted an 85% graduation rate.
"The acclaim for Joe Paterno has stemmed largely from the contrast between the high academic and moral standards he has tried to exemplify and the shameless conduct that often embarrasses and dishonors the college sport he cherishes," author Michael O'Brien wrote in a 1999 biography of Paterno, "No Ordinary Joe."
Paterno was born in 1926 in Brooklyn to second-generation Italian immigrants, according to O'Brien's book.
He attended Brown University, where he played quarterback and cornerback, according to another Penn State biography.
He coached at Penn State as an assistant from 1950 to 1965 and became head coach in 1966.
Decked out in his soon-to-become trademark thick glasses, white socks and sneakers, Paterno quickly became a memorable fixture on the football field, leading the Nittany Lions to undefeated seasons in 1968, 1969 and again in 1973 and the first national championship of his tenure in 1982.
Named National Coach of the Year five times, Paterno was added to the College Football Hall of Fame in 2006, but his induction was delayed until 2007 because of injuries he suffered in a sideline collision.
He became the winningest coach in major college football history in 2011 with 409 victories.
In addition to his exploits on the sidelines, Paterno had a significant impact on the university's academic programs.
Paterno and his wife, Suzanne, donated more than $4 million to the university over the years for faculty endowments, scholarships and building projects, according to the university.
"Penn State has been very good to both Sue and me," he said in 1998, according to his university biography.
Honored with glowing words of praise from players and presidents alike -- President Ronald Reagan said Paterno never forgot that "he is a teacher who's preparing his students not just for the season, but for life," according to a university biography -- he received the National Football Foundation and College Football Hall of Fame Distinguished American Award in 1991.
In doing so, he became the first active coach to do so, according to the biography.
"What are coaches?" he said at the dinner celebrating his award, according to his university biography. "Number one, we're teachers and we're educators. We have the same obligation as all teachers at our institutions, expect we probably have more influence over our young people than anyone other than their families," he said.
It was his perceived failure to meet those obligations that led to his downfall as the only coach many Penn State football fans had ever known.
In October, state authorities charged two university officials with misleading investigators and failing to report alleged sexual abuse in 2002, after a Penn State assistant told a grand jury he saw former assistant coach Jerry Sandusky performing what appeared to be a*** sex on a boy in a shower at the football complex.
The assistant reported it to Paterno the next day, who said he passed the report along to then-Athletic Director Tim Curley and another university executive, Gary Schultz.
Curley and Schultz left their positions shortly after the grand jury report was revealed. The next month, the university fired Paterno and Penn State President Graham Spanier.
At the time, he said in a statement released by his son, Scott Paterno, that he was "distraught" over the sex abuse scandal.
In an interview with the Washington Post published January 14, Paterno said that he felt inadequate to deal with the allegations.
"I didn't know exactly how to handle it and I was afraid to do something that might jeopardize what the university procedure was," the Post quoted him as saying. "So I backed away and turned it over to some other people, people I thought would have a little more expertise than I did. It didn't work out that way."