Wide receiver Justin Jefferson, quarterback Kirk Cousins and safety Harrison Smith have never played a preseason game under second-year coach Kevin O'Connell, and they didn't start Saturday night during the Vikings' preseason home opener at U.S. Bank Stadium, a 24-16 loss to the Tennessee Titans.

The Vikings held out 38 players against the Titans, including all 22 projected starters on the team's depth chart, following two joint practices with Tennessee this past week. The four starters who played during the preseason opener — right guard Ed Ingram, safety Camryn Bynum, nose tackle Khyiris Tonga and cornerback Akayleb Evans — joined the others on the sideline Saturday night.

Evans and rookie cornerback Mekhi Blackmon were banged up during Thursday's practice. Both are expected to make quick recoveries but were held out Saturday.

Last week at Seattle, O'Connell held out 26 players against the Seahawks, including 18 starters. Linebacker Brian Asamoah, receiver Jalen Nailor, running back Kene Nwangwu and defensive tackle Jaquelin Roy are among the young players competing for roles and roster spots who have yet to make preseason debuts due to injuries.

Roy, the fifth-round rookie, said he expects to be "full go" next week during two joint practices against the Cardinals.

A couple of healthy players — safety Josh Metellus and edge rusher Patrick Jones — may have locked up roles for this season because they didn't play against the Titans after suiting up for the preseason opener.

The Titans rested numerous starters as well, including quarterback Ryan Tannehill, running back Derrick Henry and receiver DeAndre Hopkins. Quarterback Malik Willis started for Tennessee, while rookie quarterback Will Levis suited up but did not play due to an injury suffered in Thursday's joint practice with the Vikings.

Titans receiver Mason Kinsey took a wildcat snap when Willis missed a play in the third quarter.

Strong first impression

Running back Aaron Dykes signed with the Vikings on Thursday, and the undrafted rookie out of Richmond quickly made a first impression. Dykes, a dynamic returner in college, took the opening kickoff for 38 yards. He took all five returns in the game. Rookie running back DeWayne McBride did not see any returns after handling duties in Seattle.

"Definitely a surreal moment," Dykes said. "A moment I'll never forget regardless of what happens the next couple weeks."

Cornerback NaJee Thompson, the undrafted rookie from Georgia Southern, made another play on special teams. Thompson corralled Kyle Philips in the first quarter, capping a 2-yard return for Ryan Wright's 56-yard punt.

Needs improvement

Right tackle Oli Udoh, who is projected to be the Vikings' swing tackle behind O'Neill and Christian Darrisaw, had a rough night. Udoh drew two penalties on back-to-back plays for illegal hands to the face and holding. Udoh also appeared to allow sacks on back-to-back plays in the second quarter.

A week after the Vikings were flagged nine times in Seattle, they drew 10 penalties (four declined or offsetting) on Saturday. Issues included operational hiccups: illegal formation, false start.

On defense, the Vikings were forced into burning a timeout because only 10 defenders were on the field when a substitution was botched by the Titans offense hurrying to the line. .

O'Neill could practice vs. Cardinals

Right tackle Brian O'Neill went through a pregame workout with offensive line coaches Chris Kuper and Justin Rascati on the artificial turf at U.S. Bank Stadium. O'Neill is about 8½ months removed from an avulsion fracture to his right Achilles. He sat out both joint practices and Saturday's game against the Titans, but O'Connell said last week that he would "maybe" get going next week. The Cardinals are in town Wednesday and Thursday for joint practices and the preseason finale.

Etc.

• Safety Lewis Cine, the 2022 first-round pick, started and played the entire game. After missing the tackle on a 33-yard Titans touchdown run, Cine sacked Willis near the goal line to help force a Titans field goal. "It felt good," Cine said. "But I need to be more consistent in those plays, keep them ongoing."

• Kicker Greg Joseph remained perfect this preseason. He made all four kicks against the Titans, including three field goals from 26, 33 and 45 yards. Joseph also made all three kicks in Seattle.

• Safety Theo Jackson injured the AC joint in his left shoulder, O'Connell said, during a touchdown-saving tackle near the goal line in the third quarter. Jackson was evaluated by team medical staff and returned in the fourth quarter.

• Receiver N'Keal Harry did not play after suffering a "soft-tissue" injury during warmups, per O'Connell.