Seeley's 1974 (paywalled) paper is referred to in Southwick & Moritz's
"Social Control of Air Ventilation in Colonies of Honey Bees" (J. Insect
Physiol. 33:9, 623-626, 1987;
http://citeseerx.ist.psu.edu/viewdoc/download?doi=10.1.1.502.4011&rep=rep1&type=pdf
).

Southwick & Moritz set up tiny, single-entrance boxes with 2,000 workers
plus 1 queen and observed fanning behavior and air flow. They found fanning
behavior similar to vertebrate respiration, but with a respiratory
frequency of 2.9 times per minute in the day, and longer at night. Workers
would fan to exhaust stale air. They stop fanning and heat and CO2 rises,
oxygen drops; they fan again and heat and CO2 drop, oxygen rises.

Their Figure 2 shows the neat relation between the three. CO2 varies, in
this instance, between 1.5 and 2% concentration. Yet, in the winter
cluster, atmospheric CO2 goes to 5 or 6% without fanning to remove it.
Perhaps excess fanning/roaring as seen in a queen battery box or a package
in a hot situation is a reaction mainly to heat. Maybe the bees "know" that
excess heat can lead to excess CO2 and death, and so rather than let CO2
build up to where they are too lethargic to remove it, they try to expel
the heat, even if they can't, and a positive heat feedback loop begins.
That's the danger of stopping during the day when hauling packages, or
shipping queens without proper ventilation. They don't "know" that the heat
will momentarily subside, and they do themselves no favors in expending
themselves to remove it. Correct me if I'm misunderstanding.

I've seen (very infrequently) a battery box of 100 queens with attendants
free to circulate around the cages arrive with all the attendants dead, en
route from Hawaii or California to the US midwest. Most attendants are
sticky wet, having expelled their gut contents trying to cool the "colony."
The tracking info shows they spent a couple hours in Memphis, Tennessee,
probably in the sun. In such cases, sometimes a few queens survive, located
near the vents. But they aren't in good shape. The heat feedback look might
have killed the attendants and a majority of the queens, and with their
death, the feedback loop heat source subsided and a few queens survived.
In-transit refrigeration would have made a difference in these few cases.

Tim
eastern Washington, USA

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