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Psyche 
[Vol. 92 
towards the glass rods and large numbers of workers came into 
contact with them. The high number of radiolabeled virgin queens 
resulting from all of these encounters showed that contaminated 
workers either fed, groomed, or contacted them frequently. This 
rate of distribution of radioactivity suggested that there was a very 
high level of contact between colony members. This indicates that 
all pheromones could be rapidly distributed regardless of their 
origin. This is of considerable general interest and supports hy- 
potheses of pheromonal control of behavior in social insects (Wilson 
1971). 
Radiolabel was removed from both the dead mated queens and 
live mated queens and transmitted to over 50% of the virgin queens 
within 2 h. Radiolabel was transmitted to more virgin queens from 
mated queen corpses than from live queens in two of the three trials, 
possibly because a greater number of workers have access to a dead 
queen. Seeley (1979) has shown that honeybee workers collect queen 
substance most heavily when the queen is stationary. Since the live 
queens used in our study were disturbed frequently by sampling, 
periods in which they remained stationary were limited. This could 
have reduced the number of worker-queen contacts and thus the 
quantity of radiolabel removed. Although radiolabel from the sur- 
face of live mated queens was transmitted to virgin queens at a 
slower rate, its transfer was still rapid enough to insure that it would 
have reached all of the virgin queens in less than 12 h. 
Examining the effect of food distribution on the transfer of 
radioactivity from live queens to virgin queens, we found that the 
presence of food increased the quantity of radiolabel transferred. 
Although not conclusive, our results suggested that food exchange 
may have been involved in signal transfer. We realise that these 
results could be misleading. A similar experiment with well fed 
queenless honeybee workers on 1 side of a wire mesh screen and 
starved queenright workers on the other side also showed that the 
queenright group passes “information” to the queenless group dur- 
ing transfer of food but that food and information travel in opposite 
directions (Verheijen-Voogd 1959). Also, recent research supports 
surface transport over food exchange as the primary mechanism of 
honeybee queen substance transmission (Seeley 1979). On the other 
hand, food exchange between workers has been implicated in the 
transfer of pheromone produced by immature honeybee queens 
(Free and Ferguson 1982). Because the frequency and extent of food 
