“I’m really proud of what Colton did while I was out,” Winters said. “I’m thankful for the coaches having the confidence in me to play me this week, but I’m really grateful for Colton getting us in this position, and I thought he did a great job.”

In his own, soft-spoken way, Winters was thanking Colton for the ride. But now, he’ll take the keys back, thank you very much.

On the other side of the football, the defense, the one constant all year has been the powerful, suffocating defensive line. Senior defensive tackle Josue Ortiz and Co. had held opponents to an average of just 55.4 yards per game on the ground coming into Saturday’s game.

But against Princeton, when upside down became right-side up, the run defense took the day off.

Other than one sack by Ortiz, who was pulled early in the second half after suffering a mild concussion, the usually dominant D-line got dominated. The leave-your-mouth-wide-open front seven did just that against the Tigers, but for a much different reason.

In its first five games, the Crimson D-line had allowed 277 yards on the ground.

In 60 minutes, Princeton nearly matched that number, galloping for 267 yards.

And the on-and-off secondary was off as well, surrendering nearly 300 yards in the air. The trouble really began when senior cornerback Matt Hanson had to sit out a few series after getting banged up. Without its stalwart, the secondary struggled, dropping coverages and failing to stop the relentless Princeton attack.

Eventually, after the all-Ivy senior came back in, the defensive troubles subsided, and Harvard overcame the adversity to take a far more exciting game than was previously expected.

But Murphy refused to explain away the defensive struggles by pointing to injuries to key players. And rightly so. He knows there’s serious work to be done before the level of competition picks up.

“I don’t care who goes out of a game,” Murphy said. “If Josue goes down, we have to step up. We talk about it all the time: everything we do is about adversity training ... The true test of fortitude and the true test of excellence is being able to fight through adversity. So there’s no excuses.”

—Staff writer Robert S. Samuels can be reached at robertsamuels@college.harvard.edu.