NASHVILLE, Tenn. — After fumbling a punt late in the third quarter, Cleveland Browns wide receiver Travis Benjamin was greeted at the Browns’ sideline by coach Mike Pettine.

“I said, ‘Hey, hang in there. When you have a chance to make a play, make a play.’ He didn’t go in the tank,” Pettine said.

Instead, Benjamin flushed Tennessee’s 28-3 lead with two fourth-quarter touchdown catches, capping Cleveland’s stunning comeback with his game-winner with 1:09 left as the Browns beat the Tennessee Titans 29-28 at LP Field on Sunday.

The third-year pro from Miami (Fla.) hauled in a 6-yard scoring strike from quarterback Brian Hoyer for the go-ahead touchdown, tip-toeing along the back line of the end zone to keep both feet inbounds.

“I knew I was going to high-point the ball,” Benjamin said. “We go over it in practice every day about getting two feet in.”

Benjamin’s play not only ended the Browns’ 12-game road losing streak that dated back to Dec. 1, 2012, it also represented the largest comeback by a road team in NFL history. Cleveland trailed by 25 points with 2:44 left in the first half before gradually reeling in Tennessee by dominating both sides of the scrimmage line.

Trailing 28-22, Cleveland got the ball back when backup quarterback Charlie Whitehurst was stopped on a fourth-and-inches sneak at the Titans’ 42 with 3:03 left in the game. Hoyer, who completed 21 of 37 passes for 292 yards and three scores, zipped a 23-yard completion to wide receiver Miles Austin with two minutes left.

Three plays later, Hoyer bought time until he located Benjamin with the play that finalized an incredible rally.

While the Browns (2-2) poured on the field to celebrate a potential season-turning win, the Titans (1-4) ran off the field in disbelief after eating their fourth straight loss.

The losing extended beyond the final score. Tennessee saw oft-injured starting quarterback Jake Locker (thumb) go down late in the first half and might have lost strong safety Bernard Pollard with an Achilles’ tendon injury.

What the Browns said:

“I may have to go in for an EKG test in the morning. I told the guys they’re going to have to quit doing this to me.” — Coach Mike Pettine after his team rallied from a 28-3 deficit to beat the Titans.

What the Titans said:

“It was a bit of a cheap shot, no doubt about it.” — QB Jake Locker on being hit in the head by LB Chris Kirksey after an 11-yard TD run in the second quarter.

What we learned about the Browns:

1. This team has a better offense than anyone imagined without WR Josh Gordon. They’ve scored at least 21 points in every game, mixing the running of Ben Tate with some surprising options in the passing game. WRs Travis Benjamin, Taylor Gabriel and Andrew Hawkins have joined veteran Miles Austin to make up for Gordon’s absence. By the time Gordon returns in Week 11, Cleveland could become even more dangerous.

2. First-year coach Mike Pettine might have been ripped by some for how he handled the quarterback circus in preseason, but the players are clearly buying into his message. How else can one explain comebacks from 24-0 and 28-3 deficits in the Browns’ first four games? They didn’t finish the rally from a 24-0 hole in Pittsburgh with a win, but overcoming a 25-point deficit Sunday could resonate with this team for a while as Pettine tries to build something long-lasting.

–LE Phil Taylor (knee) left the game in the first half and wasn’t able to return. Taylor had a tackle for loss. His status for next week’s home game with Pittsburgh wasn’t available immediately after the game.

–RB Ben Tate rushed 22 times for a game-high 123 yards despite having no run longer than 15 yards. Tate’s one-cut and go style works perfectly in the scheme of offensive coordinator Kyle Shanahan, and gives the offense the balance it needs to succeed. Tate is averaging 5.9 yards per carry and should flourish as he assumes the lead role in the ground game after missing time in the first three games due to injury.

–QB Brian Hoyer showed poise in the second half, rallying Cleveland from a large deficit with good decision-making and the ability to stand in the pocket in the face of a solid rush. Hoyer hit 21 of 37 for 292 yards and three scores, including the game-winning TD with 69 seconds remaining. Barring injury, it appears as though heralded rookie Johnny Manziel will continue to lug a clipboard for the season’s remainder.

What we learned about the Titans:

1. QB Jake Locker can’t be counted on to stay healthy. It’s not his fault that he hit his thumb against the helmet of CB Buster Skrine in the first half and appears to be in line to miss time again, but Locker’s injury history — he’s missed starts since 2012 with hip, shoulder, foot and wrist ailments — gives this team every reason it needs to look for another QB of the future in next spring’s draft. And it should have a high pick with which to do it, barring a shocking turnaround.

2. Tennessee’s mettle will be tested more than ever next week. A home game against winless Jacksonville won’t be easy at all, considering the Titans have lost four straight and might have to replace multiple injured starters. What’s more, after blowing a 28-3 lead Sunday, you have to wonder about this team’s mental state. Will it keep working to improve and win games or does it cash it in for the season?

–QB Jake Locker (thumb) left the game with 4:04 left in the first half after hitting it on the helmet of Cleveland CB Buster Skrine while being roughed on an incomplete pass. Locker could miss time with the injury, which would be the fifth injury that’s caused him to miss a start since becoming the starter before the 2012 season. Locker was 8 of 11 for 79 yards and a score, while rushing for 34 yards and a touchdown, before getting injured.

–SS Bernard Pollard left late in the fourth quarter with a leg injury that the team fears could be a season-ending Achilles tendon tear. Pollard was seen leaving LP Field on crutches after the team’s 29-28 loss. He had four tackles and a pass breakup against the Browns.

–CB Coty Sensabaugh (knee) might have injured his MCL in the fourth quarter of Sunday’s game. Sensabaugh actually played pretty well as a replacement for starter Blidi Wreh-Wilson, who was out with a concussion. Sensabaugh, a third-year pro out of Clemson, finished with five tackles and a pass breakup.