This actually something I pondered about this fall while inspecting thousands of hives in Southern Alberta, run by some very good beekeepers.

I noticed that some beekeepers had honey packed wall to wall in the broods and others had outside combs which were either not full, or only marginally so.  I concluded the difference was due to the strength of the flows, whether the beekeeper spreads brood in spring, the timing of supering and pulling, and perhaps, to a lesser extent, the strength and nature of the colonies themselves. 

Some beekeepers are slower to super and their bees spread out more since there is nowhere to go or put the nectar while waiting for space to be added.  Once spread wall to wall, they tend to stay that way.  If supers are added slowly, that encourages storage lower in the hive.

Hives with excluders on the second box tend to pack the brood chambers more, compared to hives without excluders or with excluders on a single.  The size of the bottom entrance may have an effect. too.

We have always kept a frame feeder in every brood chamber, first and second, and also a sheet of plastic foundation on the outside (9 and 10-frame broods).  If we were supering on time, these were seldom filled with comb, but if we were late, we saw the feeders filled with comb and the foundation drawn.  The outside position was a great place to keep a spare sheet of plastic foundation, since it was always right there if needed for brood manipulations and provided an 'overload' when things got tight.

Beekeepers report that, left to their own devices, honey bees will develop vertically before they expand horizontally.  Some books claim that this varies with lattitude and that bees tend to go more horizontally in the tropics.  I think, though, that the vertical preference in temperate regions has mostly to do with heat conservation, since heat rises and if the bees work vertically, they get more use from each calory before it escapes upwards. 

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