Thanks for the posts Pete!

Allow me to add some of my own practical observations, from one who always
uses smoke, but sparingly.

In general, the only bees that need to be smoked are those on the periphery
of the cluster, esp near any entrances.  That is where bees acting as
guards tend to be found.

By closely observing the behavior of those guards, one can determine
exactly the amount of smoke required.  The guards are the only bees that
pay any attention to the beekeeper.  They are the bees that watch you and
follow your movements.

Any bee watching you is aware of your presence.   If I see eyes watching me
from the interspaces between the combs,  I apply just enough smoke to turn
those bees around, so that they no longer face me.  So long as no bees are
facing me (and are thus aware of my presence), I can handle the frames with
impunity.

I do not discount the effect of smoke on the diminution of the olfactory
receptors to alarm pheromone, but in general, the effect of smoke, as I use
it, is exhibited prior to any release of alarm pheromone.  A small amount
of smoke changes the behavior of the guards, gently repels bees from the
tops of the combs, and if applied to an open comb, causes the younger
workers to move off the open brood and to engorge on nectar.

Many beekeepers, IMHO, apply far more smoke than necessary.

As far as the effect of smoke upon olfaction.  It is only after I have make
a serious handling mistake that alarm pheromone is released.  I can smell
it, and I can easily see how it elicits defensive response in any older
workers that happen to be in the odor plume (the guards take their stances,
start moving rapidly, and give warning bumps).  If I persist at that moment
in adding to the triggers for release of alarm pheromone (my own rapid
movements, vibration, etc), additional workers will release pheromone, and
the colony defense response quickly grows.

On the other hand, if I simply give the bees a moment for the pheromone to
dissipate (which can be determined by watching their behavior), and then
apply a small amount of smoke to get them to take their eyes off me, I can
resume working.

However, once a full defensive response is initiated (as in the case of
Africanized bees), the beekeeper has the choice of either ceasing working
that hive (or perhaps the entire apiary), or stepping up his use of smoke
to an extreme level.  At that point, the olfactory suppression likely comes
into play.

-- 
Randy Oliver
Grass Valley, CA
www.ScientificBeekeeping.com

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