Michigan can forget a perfect run through the Big Ten.
The Wolverines' flaws were unspooled one missed field goal, one costly interception at a time.
Bill Belton ran for a 2-yard touchdown in the fourth overtime to lift Penn State to a 43-40 victory over No. 18 Michigan in maybe the biggest win in coach Bill O'Brien's two seasons with the Nittany Lions.
Penn State's Christian Hackenberg was sensational in regulation on the tying drive, completing passes of 29 yards to Brandon Felder and 33 yards to Allen Robinson to bring the ball to the 1 with 29 seconds left.
The freshman quarterback sneaked in for the first rushing TD against Michigan this season, tying the game at 34-all.
Brendan Gibbons kicked two field goals and missed two — one was blocked — in OT for the Wolverines (5-1, 1-1). Sam Ficken kicked two, but passed on even trying another on the last possession. Instead, Penn State went on fourth-and-1 from the 16 and Belton dove ahead for 3 yards. Three plays later, he was in the end zone, and the party was on.
Just not for Michigan.
"I think you always work hard at any time, but to lose in overtime, if you let it take a toll, it will," coach Brady Hoke said.
There were plenty of blown chances for the Wolverines. Gibbons missed 3 of 7 field goals, Devin Gardner threw two interceptions and lost a fumble, and a pass interference penalty aided Penn State's winning drive.
"We had all kinds of opportunities at every position," Hoke said.
The Nittany Lions (4-2, 1-1 Big Ten) stormed the field in wild celebration, sending more than 100,000 fans into a frenzy.
"I just ran somewhere and hit somebody," Hackenberg said. "I saw him bounce it and I saw he had the edge and after that I just ran over to the sideline to celebrate with my teammates.
Still recovering from scandal, the Nittany Lions gave 107,884 fans at Beaver Stadium at long last a milestone celebration in Happy Valley. The Nittany Lions mobbed O'Brien, and he led the charge toward the students in the stands to sing the fight song. Penn State's victory bell rang long after the players had stunned the unbeaten Wolverines.
"Nothing should amaze you," O'Brien said. "There's going to be twists and turns. These are tough kids. They love Penn State. They love playing for each other. The locker room is such a great scene right now because these kids really believe in each other."
Hackenberg, who threw for 305 yards, made it possible with plenty of help from his receivers on the final drive of the fourth.
Robinson expanded his 6-foot-3 frame about another 4 inches on a fantastic, outstretched leap in the corner that sent him crashing down a yard from the goal line.
"It was just me and (the defender) over there and it was a 50-50 ball," he said. "He threw it up and gave me an opportunity to make the catch and I was able to make the play."
Hackenberg plowed through for one sweet yard on the next play.
The 18-year-old freshman threw three touchdown passes in the first half to stake Penn State to a 21-10 lead.
Gardner put the Wolverines ahead with touchdown passes in the third and fourth quarters.
Gardner connected with Jeremy Gallon for a 16-yard TD pass for a 27-24 lead with 28 seconds left in the third. Gardner was 15 of 28 for three touchdowns and he rushed for 121 yards.
He then hit Devin Funchess in the back of the end zone for a 37-yard TD and a 34-24 lead. Funchess had 112 yards receiving and two scores.
Ficken hit a 43-yarder with 6:35 left to make it 34-27, keeping Hackenberg and the Nittany Lions alive for one thrilling finish.
Gibbons missed a 52-yarder with 2 seconds left in regulation.
Michigan defensive end Frank Clark returned a fumble for a score in the third quarter to cut it to 21-17, and spark Michigan's comeback.
Michigan did get a boost with the return of star linebacker Jake Ryan, who had yet to play this season after he suffered a torn anterior cruciate ligament in the spring.
"It's not about mistakes. It's about us executing," Ryan said. "On some plays we didn't do that and that's what we need to get done. That's what we need to get fixed."
Michigan offensive lineman Taylor Lewan, an All-American who decided to pass on the NFL and return for his senior season, sat out the second half with an undisclosed injury.
We didn't take advantage of the opportunities we had," Ryan said. "We just have to keep our heads up and move on to next week. That's Indiana."