EAST LANSING, Mich. (AP) Michigan State has to replace its star quarterback, its top receivers and some key offensive linemen from 2015.
That leaves the Spartans' group of running backs - which returns largely intact - under a decent amount of pressure.
''The running backs, we're the ones that's considered as veterans on the team,'' Madre London said. ''We all played.''
LJ Scott, Gerald Holmes and London combined for 1,739 yards on 375 carries last season, and all three of those tailbacks return. When quarterback Connor Cook was limited by injury issues late in the season, the running game pushed Michigan State to crucial victories over Ohio State and Iowa.
Scott led the team with 146 carries for 699 yards. Holmes (110) and London (119) had about the same number of carries, in part because the latter missed some time with an ankle injury.
Holmes began to play a bigger role during the second half of the season. Starting on Oct. 24 against Indiana, he had a touchdown in five straight games - his first major contribution after redshirting in 2013 and playing sparingly the following year.
''Obviously, redshirting my freshman year, and being out of the mix for a while ... being able to get back on the field was always a blessing, and it felt good,'' he said. ''I was pretty surprised, the progress I made.''
Michigan State went 12-2, losing to Alabama after reaching the College Football Playoff. It owed a lot of its success on offense not just to Cook and receivers Aaron Burbridge and Macgarrett Kings - all seniors - but also to a talented offensive line that overcame a number of injuries.
The Spartans will have to replace linemen Jack Allen, Donavon Clark and Jack Conklin. Allen and Clark were fifth-year seniors, and Conklin is skipping his senior season to go to the NFL draft.
''That's part of the game,'' London said. ''By us being a great program that we are right now, we're just going to fill in the blanks.''
Last season, it was the running backs who were the unproven commodities. Jeremy Langford and Nick Hill had been the team's top rushers in 2014, and with them gone, it was time for Scott and London to take over. Holmes emerged later in the season.
They shared the carries to such a degree that Scott, London and Holmes each had only one game with at least 100 yards rushing. Michigan State had enough depth at running back that Delton Williams wasn't much of a factor, carrying the ball only 26 times.
Williams was suspended last March, shortly after being arraigned on a charge of brandishing a firearm in public. He later pleaded guilty to a reduced charge. Williams was allowed to return to the Spartans in the fall as a non-scholarship athlete and the 6-foot-1, 230-pounder is listed as a fullback on the depth chart.
''He's making the transition right now, and again, it's a slow process,'' coach Mark Dantonio said. ''Still can play tailback and still does at times.''
Dantonio has said Williams will go back on scholarship in the summer. Now entering his senior season, Williams said he tries to be a good mentor to younger players, hoping they won't have to go through anything similar to his off-field problems.
''The stuff that I went through last year, it all molded me to be who I am today. I'm not mad about anything that I went through,'' he said. ''The consequences and stuff I had to go through, I really appreciate Coach `D' giving me another chance to be here.''
Follow Noah Trister at www.Twitter.com/noahtrister