Yes, but how often are main line services crush loaded, apart from at their London termini when dwell time doesn't matter, and at how many places do they have a substantial crush load interchange on and off, like the Jubilee Line at Waterloo?
Regularly crush loaded. And with 2 doors per coach (and narrow doors on the 455s at that), it only needs one way flow fir there to be long dwells. For big two way flows, off the top of my head:
Clapham J, Wimbledon, Richmond, Balham, E Croydon, Denmark Hill, Lewisham, and London Bridge (6-9).
Notably the Elizabeth line has 60 second dwells throughout the core, even with 3 sets of door per coach side, and they are often needed.