The anti-waggle dance: use of the stop signal as negative feedback

Abstract

Numerous activities within honey bee (Apis mellifera L.) colonies rely on feedback loops for organization at the group level. Classic examples of these self-organizing behaviors occur during foraging and swarm nest site selection. The waggle dance provides positive feedback, promoting foraging at a specific location or increased scouting at a potential nest site. Rather less well known than the waggle dance is the stop signal, a short vibration often delivered while butting against a dancing bee. It is currently best understood as a counter to the waggle dance, offering negative feedback toward the advertised foraging location or nest site. When the stop signal is received by a waggle dancer she is more likely to terminate her dance early and retire from the dance floor. Bees that experienced danger or overcrowding at a food source are more likely to perform the stop signal upon their return to the colony, resulting in an inhibition of foraging at that location. During a swarm's nest site selection process, scout bees that visited a different site than the one being advertised are more likely to stop-signal the waggle dancer than are scouts that had visited the same site. Over time, the scout bees build recruitment to a single site until a quorum is reached and the swarm can move to it. The balance between the positive feedback from the waggle dance and the negative feedback from the stop signal allows for a more sensitive adjustment of response from the colony as a unit. Many of the processes associated with the feedback loops organizing a honey bee colony's activities are in striking parallel to other systems, such as intercellular interactions involved in motor neuron function.

Long-term trends in the honeybee ‘whooping signal’ revealed by automated detection

“…Producing a modulatory signal makes perfect functional sense under the condition of a colony experiencing an excessive surge in its nectar influx, so that the colony needs to supress the recruitment of additional nectar foragers and facilitate the recruitment of additional nectar receivers [ 13 ]. Inhibiting waggle dancers can also have a secondary effect in that more nectar receivers can be recruited (reviewed in Keitzman and Visscher [ 23 ]). Thus it is not surprising to see that tremble dancers act to inhibit waggle dancers, as reported by Lau and Nieh [ 9 ], Nieh [ 7 ] and Thom et al .…”

Honey Bee Inhibitory Signaling Is Tuned to Threat Severity and Can Act as a Colony Alarm Signal

“…mellifera foragers produce stop signals to inhibit waggle dancing [ 19 – 21 ] for dangerous food sources. A stop signal is a 300–400 Hz vibrational signal with a duration of approximately 150 ms [ 20 , 22 ] that a worker usually delivers while butting its head into the body of the receiver, causing the receiver to momentarily freeze [ 12 , 19 , 23 , 24 ]. Attacks by wasp and spider predators [ 12 ] and conspecifics [ 18 ] elicit stop signals, which inhibit waggle dancing.…”

The Waggle Dance as an Intended Flight: A Cognitive Perspective

“…Recruits that had trophallactic interactions with the dancer expect the same quality of the food source (odor, taste) when arriving there [1]. Stop signals performed by following bees are considered as a feedback to the dancer, which requires some form of expectation about the potential outcome if the receiver would apply the dance-indicated behavior [21,[45][46][47][48][49]. Stop signals produced are particularly informative in our context when the dancer advertises a food source that is dangerous or of inferior quality [46,49].…”