MINNEAPOLIS - Defensive veterans like Dan Connor and Donnie Johnson spoke this past week about having to stop Minnesota's always-potent running game. Penn State's defense accomplished that.

It did not, however, manage to contain the Golden Gophers through the air.

Minnesota quarterback Bryan Cupito was 25-of-36 and threw for two touchdowns. His 347 yards tie him with Billy Dicken of Purdue (1997) for the 15th-highest total ever put up against Penn State and the most since Jeff Smoker lit up the Nittany Lions for 357 yards in 2003.

Slants and other quick routes proved difficult for Penn State to defend, as the defensive line often reached the backfield, but not in time to reach Cupito before he fired to an open receiver.

Cupito's two favorite targets, tight end Matt Spaeth and receiver Logan Payne, each grabbed six catches for 99 and 94 yards, respectively.

In the first quarter, with the ball at Penn State's 30, Sean Lee came on a blitz. Spaeth ran a post pattern on which he went to the outside of linebacker Paul Posluszny, then shifted to the inside of safety Anthony Scirrotto, freeing himself for Cupito's throw, then running unimpeded into the end zone for the game's first score.

"Defensively, we obviously didn't cover the tight end like we should have," Posluszny said. "We missed some assignments and that led to big plays."

With 2:10 to play in regulation and down 21-14, Cupito hit Payne on a slant with the game on the line on fourth-and-2.

"I knew I was going to be open with the way they were playing me, and the way they were playing that route I knew I was going to have a chance to make a big play," said Payne, who made the catch and eluded a handful of Penn State defenders on his way to a 42-yard gain that helped set up the tying touchdown.

A big problem for Penn State's secondary was poor tackling, which allowed moderate gains to become significant.

"We just needed to be making simple tackles," defensive tackle Jay Alford said. "That's the biggest thing. If we make the tackles, they don't get first downs. That's pretty much it."

On the bright side, Minnesota was held to its lowest rushing total since a Nov. 6, 2004 loss to Wisconsin. The Golden Gophers amassed merely 68 net yards on the day, averaging two yards per attempt.

Alford led the way, spending a good amount of time in Minnesota's backfield. He finished with 1.5 sacks, four tackles for a loss and seven total tackles.

"This week he was excited and ready to go," Ed Johnson said of his battery-mate. "He had a lot of momentum today; I was down a little bit and he did a good job of keeping me up."

Josh Gaines, filling in for Jim Shaw at defensive end, also played well, combining with Alford on an early sack of Cupito.

With No. 4 Michigan coming into Beaver Stadium Saturday, the defense will look to get stronger in a hurry against the quick routes, as the Wolverines' have historically taken a liking to slant patterns.

The general feeling among the Lions defensive backs and linebackers, though, was that they were capable of putting a better product on the field than what was shown against Minnesota's passing game.

"It shouldn't have gone down to the wire like that -- let's put it that way," Posluszny said.