That's an interesting paper and worth reading. Not sure it has answered my question but I did learn some things from it that I had not considered.

This was the quote I resonated with :-)

"... the discoveries we highlight here only capture a small fraction of the mechanisms of the model."

It sometimes feels a bit like papers on cellular biology with DNA discussions in which descriptions of the enzymes and proteins involved are insightful but the mechanism that operates the reaction remains opaque.