At Purdue, where quarterbacks have largely been the storyline through its history (especially when the program has been successful), it’s noteworthy when the defense takes the headlines.

But at Saturday’s spring game, the Boilermakers’ defense won in a modified scoring system, getting a late takeaway for a 45-42 win. Perhaps the defense, which was a surprise last season, can continue to be a bright spot for Purdue, which will enter 2022 with hopes of making a Big Ten West run. Or maybe the offense, led by quarterback Aiden O’Connell, can get healthy in the offseason, see the return of several play-makers, and become a feared unit in the league.

Whatever happens, though, Purdue leaves the spring with plenty of questions. Below are several:

Will the line hamper everything?

Purdue’s offensive line isn’t set — not even close — as it enters the offseason, and that was evident during Saturday’s game.

It had moments, but not enough of them. Yet it’s clear that the group that started Saturday, which included Cam Craig at left tackle, Spencer Holstege at left guard, Gus Hartwig at center, Jared Bycznski at right guard and Marcus Mbow at right tackle, will not be the same as the one that starts Game 1 vs. Penn State this fall. Only Holstege and Hartwig are likely to stay. Eric Miller, a left tackle, was out, as was Florida International guard Sione Finau. Add those 2 back into the mix as likely starters and Purdue’s line gets not only better, but deeper.

It’s imperative for the Boilermakers to develop a competent line, because O’Connell is a pocket passer, who while able to navigate pressure in the pocket isn’t likely to be able to escape it with his feet.

Is the D as good as thought?

Purdue’s defense kept the offense off the scoreboard for the Boilermakers’ first 9 drives on Saturday, rolling up a 28-0 lead by halftime.

It was an impressive display, when also considering the defense tallied 2 interceptions as well, plus had a strip-sack in the 4th quarter that sealed the win.

But is it real? Purdue had many of its skill position players out for the spring game, including arguably 4 of its top 5 receivers (only Tyrone Tracy, the Iowa transfer, was available among expected the top-line group), was missing a couple offensive line and had tight ends who had been banged up much of the spring. Still, Jeff Brohm feels good about his defense, particularly on the D-line, where he thinks the Boilermakers are at least 2-deep.

Defensive tackle Branson Deen has a chance to be a star, as does fellow interior lineman Lawrence Johnson. Kydron Jenkins could continue to develop at Leo, the hybrid linebacker/end position, after a breakout performance there as the backup last season. And Jack Sullivan is a versatile player who can line up on the inside or out, like he did in starting for George Karlaftis in the Music City Bowl, after the All-American had decided to enter the draft early.

Is there a pass rush?

However …

With all there is to like about Purdue’s defensive line, a question remains: Can Purdue generate a pass rush, now that Karlaftis and DeMarcus Mitchell are departed?

On Saturday, Purdue got pressure — but was it a result of a hodgepodge offensive line — and collected 2 sacks, 1 each from Khordae Sydnor and Yanni Karlaftis. But the Boilermaker will want more productivity in the fall. Maybe Purdue will get a boost from the continued development of freshman Joe Sullivan, who was an early enrollee at Purdue and participated in the spring, or perhaps fellow freshman Nic Caraway, a 4-star prospect, can make a Day 1 splash in the fall. Or maybe rather than having a No. 1 pass rush, one who defenses had to game plan for every Saturday, as was the case with George Karlaftis, the opponent will need to try to account for multiple rushers on the Boilermakers’ front.

Whatever the scenario, it’s a question to watch — perhaps the question for the defense — headed to the fall.

Will there be any running game?

Purdue has finished last in the Big Ten in rushing the last 2 seasons.

And, as it leaves the spring, there’s not much reason to think that’ll change in 2022.

The Boilermakers have only a couple of experienced backs in returning starter King Doerue and transfer Sampson James, who arrived from Indiana during training camp last season, then had to sit out the year. But while both can be effective between the tackles, neither has game-breaking type of speed to the perimeter, and that’s what the Boilermakers lack.

Walk-on running back Caleb Lahey led Purdue in the spring game with 13 carries for 61 yards and a TD.

Brohm is continuing to search the transfer portal for help in the backfield, having missed out on a couple possibilities in the winter, but maybe now that teams are wrapping up their spring drills other options will emerge.

Can Purdue accept success?

Purdue won 9 games last season, the most since its 2003.

And the Boilermakers are coming off a dramatic victory over SEC rival Tennessee in the Music City Bowl, leaving many to wonder whether they can take another step forward this season, perhaps even be a dark horse to compete in the Big Ten West.

Whether they can (or can’t) might depend on how well they adapt to success. It’s unlikely, for instance, that Purdue catches Penn State off guard in the season-opener in September. But if the Boilermakers can get by the Nittany Lions, it’s conceivable that they’ll be favorites in 5 of the following 6 games. That’s not how Purdue ran its record to 9-4 last season; rather it did so by upsetting 2 top-5 ranked foes to get a jump-start for the second half of the year.

Now, the Boilermakers will have a chance to get out of the gate hot.