Not sure about the 400F, but on a 750, the right side axle spacer is a part of the speedo drive. So, when you clamp the axle in place, it presses the speedo drive against the right side wheel bearing's inner race, which itself presses against the wheel's center spacer. So, when you clamp down the axle nut, the speedo housing becomes clamped to the axle and both will spin as a unit, but it you hold the wheel by the axle and keep it from spinning (like when you clamp it into the forks) then the speedo drive will not spin either.
mystic_1