LINCOLN, Neb. — As time expired, Austen Arnaud sprinted toward mid-field until he found the person he was looking for.

“I’m proud of you; I’m proud of you,” Arnaud screamed as he jumped into freshman quarterback Jerome Tiller’s arms.

All Tiller could do was stand there with a smile, soaking in the aftermath of his first career start, the Cyclones’ eight forced turnovers and his team’s 9-7 win against Nebraska in Memorial Stadium — the program’s first since 1977.

“You come into a stadium like this, and you play without the Big 12’s leading rusher [Alexander Robinson], you play without your starting quarterback [Arnaud] — not to mention a lengthy list of individual players who were sick,” said coach Paul Rhoads, who was visibly emotional after the game, “and to extend that kind of effort is just a fabulous, phenomenal effort, and I couldn’t be prouder of our football team.”

The Cyclones relied on an opportunistic defense, which, with the help of the eight turnovers, held Nebraska to seven points — the Huskers’ lowest total against Iowa State since 1959.

“We came into this game supremely confident with our defense,” said defensive tackle Nate Frere. “We really showed up when we needed to. There was a few times when they were knocking on the door and a guy ripped the ball out. That’s the kind of things you need to win these big games.”

Nebraska brought the ball to within striking distance on numerous occasions, but four turnovers within 10 yards of the end zone, including a fumble on the Cyclones’ 5-yard line late in the third quarter by Dontrayevous Robinson, ultimately doomed the Huskers.

With Arnaud nursing a bruised throwing hand and Robinson still feeling the lingering effects of a groin injury, Iowa State turned to Tiller and backup running back Jeremiah Schwartz to help guide the offense.

“If you picked the perfect place for your backup quarterback to start a game, I promise you it wouldn’t be Memorial Stadium,” Rhoads said after the game.

Despite the hostile environment, Tiller remained poised under center.

He absorbed the heavy pressure he received from the Nebraska defense, throwing the ball away safely when needed. He finished the day 9-of-19 for 102 yards, and added another 65 yards on the ground.

“The most important thing he did today was not turning the ball over, and that gave us a chance. He did a great job in there today,” said center Reggie Stephens.

Iowa State’s defining offensive moment came in the second quarter, following punter Mike Brandtner’s 20-yard sideline dash on a punt fake — a 47-yard touchdown pass from Tiller to Jake Williams.

With the Cyclones facing fourth and five on their own 33-yard line, Iowa State made the call for the fake.

“There are a lot times when teams are looking for that play, but it was a perfect time to run that play,” Brandtner said. “No one rushed, and I saw red jerseys turn to go downfield, and I just took off down the sidelines.”

After taking a timeout to regroup, Tiller and the offense wasted no time to take a shot at the reeling Husker defense.

On the first play, Tiller let off a long, arching pass toward the end zone.

“I let the ball go and I saw Jake and thought, ‘this is a touchdown,’” Tiller said.

Williams fought off his lone defender and, as he did against Kansas State two weeks ago with his last-second touchdown grab, came up with Iowa State’s signature play of the game.

Williams’ touchdown put the Cyclones up 9-7, which proved to be enough.

Nebraska had two opportunities in the last four minutes to put together at least a go-ahead field goal, but the Cyclone defense held, ending both drives with an interception.

Following the final interception by middle linebacker Jesse Smith, Tiller took the final snap of the game and placed his knee to the ground. The final seconds of the game melted away and the celebration among the Cyclones ensued.

“When that clock stopped, my stomach dropped; I was so excited,” Tiller said.

Besides snapping a 14-game Big 12 road losing streak, Iowa State finds itself one game from bowl eligibly and securing its first non-losing season since 2005. The Cyclones are now also in a position to compete for a Big 12 North division title, which would earn them a spot in the Big 12 campionship game.

“Our goal and expectation for this team this year is to improve, and I think we’ve done that,” Rhoads said. “The result has led us to five victories and a team that feels pretty confident with the direction of this program.”

But in the moments following their upset victory in front of a shocked Husker crowd, the Cyclones reveled in a feeling Iowa State hadn’t experienced in Nebraska for 32 years.

“It feels absolutely unbelievable. I don’t think I’m going to stop smiling until we get back to Ames,” Williams said.