Mainshaft is supported in front input shaft in a pilot bearing. When the bearing suffers a 'critical incident' (perhaps hardening starts to let go) and starts to wear badly, gear running clearances open up and the gears start to whine and wear accelerates. Similarly, layshaft bearing wear and fifth (overdrive) gear whine are associated. But it sounds like your main drama is in the middle of the box, with third and fourth noisy.
Now, #3 son's Defender R380 has just been replaced; but before that, did 84000 km with the howl getting louder - not intrusive, but noticeable. And it has been mostly well laden, all over Kimberley, sand, mud, you name it.
Perhaps he was lucky - it held together.
Telltale signs 1. increasing noise, both bearing grumble and gear howl. 2. Unusual slime in gear oil, sort of silvery!! 3. Radio volume not enough to cover the problem. He had it all !
Check out the colour of the gearbox oil - and any contaminants therein.
Fix = out with the box and rebuild it.
HTH and good luck :-)
Pierre
