ALAMEDA (AP) — The loss of Justin Tuck to a season-ending injury and Aldon Smith to a yearlong suspension figured to be a big blow to the Oakland Raiders’ pass rush.

Instead, it had had the opposite effect with the maturation Khalil Mack, the NFL sacks leader who has been dominant over the past month and is coming off a five-sack afternoon against the Denver Broncos.

The rapid development of defensive ends Denico Autry and Mario Edwards Jr. has been just as critical.

Edwards, the 35th overall pick in the draft, has been stellar over the past month and is second among Oakland defensive linemen in tackles.

Autry also has played better since Tuck and Smith were lost, and is coming off his best game of the season when he had two hits and five hurries on Denver quarterback Brock Osweiler.

They might not be Tuck and Smith but the duo is definitely making an impact in Oakland.

“It’s just the knowledge that those two old-heads dropped on us,” Autry said Thursday. “We know what we have to do. It was up to us now.”

Autry is an undrafted free agent who spent part of 2014 on the practice squad and played sparingly as a backup. He went into training camp this year buried on the depth chart but worked himself into the defensive line rotation.

Tuck’s season-ending pectoral injury created an opening up front and Autry — who also gets work in at defensive tackle — took over.

So far, it’s been a good fit. Autry has 22 tackles and two sacks, and has used his size and long arms to knock down a team-leading three passes at the line of scrimmage. The 6-foot-5-inch defensive lineman also blocked one field goal and one extra point this season.

“(He’s) consistently playing hard, consistently understanding where he’s supposed to be, consistently understanding It’s a team defense,” Raiders defensive coordinator Ken Norton Jr. said. “If they all work hard and work together, there’s enough sacks and pressures to go around for everybody.”

Edwards, Oakland’s second-round draft pick, has been just as effective after moving into the starting lineup in early October. The rookie defensive end had breakout game with 11 tackles and a sack against Minnesota on Nov. 15. He’s backed that up with four consecutive solid performances including last week’s win over Denver when Edwards made five tackles and forced a fumble.

Oakland nose tackle Dan Williams, who signed with the Raiders as a free agent this offseason after spending his first five seasons in Arizona, isn’t surprised.

“We just started to jell better as the season went on,” Williams said. “Also, guys have put in a lot of time in studying, identifying protections and we’ve just been playing off of each other. One of the half-sacks I got was coming off of Denico. He was getting pressure and I just came around. Everybody’s playing off of each other and we’re getting more familiar, and in the end, we’re getting results.”

Mack is a potential defensive player of the year candidate and remains the key to Oakland’s defense. In the four games since Smith was suspended for a full calendar year by NFL Commissioner Roger Goodell, Mack has 29 tackles, nine sacks, four stops behind the line of scrimmage and a forced fumble.

But as Raiders coach Jack Del Rio pointed out earlier in the week, Mack’s big day against Denver came as a result of what Oakland’s defensive line as a whole was doing.

“On a couple of occasions it’s the pressure that Mario’s getting, the pressure that Denico’s getting that doesn’t allow the quarterback a step-up lane, and now Khalil is able to get the sacks,” Del Rio said. “A lot of good effort up front, and obviously Khalil ended up finishing with the sack numbers, but some good production out of that group.”

Autry said he and Williams had no choice but to step up once Tuck was hurt and Smith was suspended.

“We are held to high standards so we have to fill the shoes of the guys who were before us,” Autry said. “That’s understood. If we don’t do it, I’m pretty sure they’ll get somebody in here who will.”

Notes: RT Austin Howard missed a second straight practice with a knee injury. Starting right guard J’Marcus Webb practiced at tackle while rookie Jon Feliciano took reps at right guard with the first-team offense. ... Mack (knee) was limited in practice after being held out a day earlier. . Safety Charles Woodson (shoulder), WR Amari Cooper (foot) and RB/KR Taiwan Jones (knee) were also limited. . Safety Nate Allen (knee) did not practice after being limited on Wednesday.