Carpenter bee thorax vibration and force generation inform pollen release mechanisms during floral buzzing.

Approximately 10% of flowering plant species conceal their pollen within tube-like poricidal anthers. Bees extract pollen from poricidal anthers via floral buzzing, a behavior during which they apply cyclic forces by biting the anther and rapidly contracting their flight muscles. The success of pollen extraction during floral buzzing relies on the direction and magnitude of the forces applied by the bees, yet these forces and forcing directions have not been previously quantified. In this work, we developed an experiment to simultaneously measure the directional forces and thorax kinematics produced by carpenter bees (Xylocopa californica) during defensive buzzing, a behavior regulated by similar physiological mechanisms as floral buzzing. We found that the buzzing frequencies averaged about 130 Hz and were highly variable within individuals. Force amplitudes were on average 170 mN, but at times reached nearly 500 mN. These forces were 30-80 times greater than the weight of the bees tested. The two largest forces occurred within a plane formed by the bees' flight muscles. Force amplitudes were moderately correlated with thorax displacement, velocity and acceleration amplitudes but only weakly correlated with buzzing frequency. Linear models developed through this work provide a mechanism to estimate forces produced during non-flight behaviors based on thorax kinematic measurements in carpenter bees. Based on the buzzing frequencies, individual bee's capacity to vary buzz frequency and predominant forcing directions, we hypothesize that carpenter bees leverage vibration amplification to increase the deformation of poricidal anthers, and hence the amount of pollen ejected.

Slumber in a cell: honeycomb used by honey bees for food, brood, heating and sleeping.

Sleep appears to play an important role in the lives of honey bees, but to understand how and why, it is essential to accurately identify sleep, and to know when and where it occurs. Viewing normally obscured honey bees in their nests would be necessary to calculate the total quantity and quality of sleep and sleep's relevance to the health and dynamics of a honey bee and its colony. Western honey bees (Apis mellifera) spend much of their time inside cells, and are visible only by the tips of their abdomens when viewed through the walls of an observation hive, or on frames pulled from a typical beehive. Prior studies have suggested that honey bees spend some of their time inside cells resting or sleeping, with ventilatory movements of the abdomen serving as a telltale sign distinguishing sleep from other behaviors. Bouts of abdominal pulses broken by extended pauses (discontinuous ventilation) in an otherwise relatively immobile bee appears to indicate sleep. Can viewing the tips of abdomens consistently and predictably indicate what is happening with the rest of a bee's body when inserted deep inside a honeycomb cell? To distinguish a sleeping bee from a bee maintaining cells, eating, or heating developing brood, we used a miniature observation hive with slices of honeycomb turned in cross-section, and filmed the exposed cells with an infrared-sensitive video camera and a thermal camera. Thermal imaging helped us identify heating bees, but simply observing ventilatory movements, as well as larger motions of the posterior tip of a bee's abdomen was sufficient to noninvasively and predictably distinguish heating and sleeping inside comb cells. Neither behavior is associated with large motions of the abdomen, but heating demands continuous (vs. discontinuous) ventilatory pulsing. Among the four behaviors observed inside cells, sleeping constituted 16.9% of observations. Accuracy of identifying sleep when restricted to viewing only the tip of an abdomen was 86.6%, and heating was 73.0%. Monitoring abdominal movements of honey bees offers anyone with a view of honeycomb the ability to more fully monitor when and where behaviors of interest are exhibited in a bustling nest.