Ultimately, neither he nor Spanier could withstand the backlash that started when longtime defensive coordinator Jerry Sandusky was charged Saturday with molesting eight boys, and two administrators were accused of covering up one of his assaults.
Paterno and Spanier were not charged, but became targets of relentless criticism that they ignored or failed to act on troubling signs about Sandusky's conduct.
Board of Trustees vice chairman John P. Surma said university provost Rodney Erickson would take over as acting president. Tom Bradley, the football team's defensive coordinator, was named interim head coach.
"These decisions were made after careful deliberations and in the best interest of the university as a whole," Surma said at a late-night news briefing at a hotel following the board's third meeting in four nights.
The news about Paterno drew gasps from the crowd and instantly rippled across the campus, where students gathered in swelling groups.
Thousands took to the streets of downtown State College after the trustees' news briefing. They were met by police in riot gear, who fired tear gas to disperse the crowds. No violence was reported.
Students climbed trees and stood on rooftops along Beaver Avenue, chanting "We want Jo Pa!" and "One more game!" Some carried signs reading "We (heart) Joe Pa."
The drama came at the end of a day when the coach appeared to be fighting to stay through the end of the season - a battle he lost.
"Right now I'm not the coach," Paterno told reporters clustered outside his house late Wednesday. "And I've got to get used to that."
