Now I'm wiser. The V1 in the rack is in deed the input valve, or one half of it (one triode) is - as is the case in any Fender or Marshall or what-have-you. The other half of it in an ordinary amp would have been used for the first gain stage, if there was any, or in the case of e.g. vintage Marshals taken care of the second input.
This puts us in a sort of dilemma: In a Fender tweed of Marshall we want this to be a harmonically rich and warm, maybe even dark valve - like the Mullard, Tung-sol or other. In a Fender blackface and similar we would want a 7025; a bright valve for a twangier tone.
The V2 in the amp... probably it's just a buffer of sorts. It doesn't appear to do any inverting. The triodes are not coupled similarly in parallell as would be the case with inverters. Also any inverting should be done just before the powervalves, no earlier than that.