Resilience of honeybee colonies via common stomach: A model of self-regulation of foraging

Abstract

We propose a new regulation mechanism based on the idea of the “common stomach” to explain several aspects of the resilience and homeostatic regulation of honeybee colonies.This mechanism exploits shared pools of substances (pollen, nectar, workers, brood) that modulate recruitment, abandonment and allocation patterns at the colony-level and enable bees to perform several survival strategies to cope with difficult circumstances: Lack of proteins leads to reduced feeding of young brood, to early capping of old brood and to regaining of already spent proteins through brood cannibalism. We modeled this system by linear interaction terms and mass-action law. To test the predictive power of the model of this regulatory mechanism we compared our model predictions to experimental data of several studies. These comparisons show that the proposed regulation mechanism can explain a variety of colony level behaviors. Detailed analysis of the model revealed that these mechanisms could explain the resilience, stability and self-regulation observed in honeybee colonies. We found that manipulation of material flow and applying sudden perturbations to colony stocks are quickly compensated by a resulting counter-acting shift in task selection. Selective analysis of feedback loops allowed us to discriminate the importance of different feedback loops in self-regulation of honeybee colonies. We stress that a network of simple proximate mechanisms can explain significant colony-level abilities that can also be seen as ultimate reasoning of the evolutionary trajectory of honeybees.

Integral feedback control is at the core of task allocation and resilience of insect societies

“…The balance between these forager groups is regulated by the protein saturation of worker bees that act as a common protein stomach. This regulation system can offset detrimental perturbation, such as rainy periods, pollen traps, or loss of foragers (23).…”

“…We studied the task allocation mechanisms of wasps (17–20), ants (21), and honey bees (22, 23), and in all cases, we observed that workers were able to switch tasks without requiring noticeable changes in development or learning experience, or that these switches mostly caused short-term time delays (24). These switches result in rapid adjustment of the workforce, allowing the colony to compensate for disturbances.…”

Solitary bee larvae prioritize carbohydrate over protein in parentally provided pollen

“…Helm et al, 2017); and researchers have often inferred offspring requirements indirectly from the patterns of alloparental feeding within studies more broadly focused on adult foraging (see Cook et al., 2010; Dussutour & Simpson, 2009; Vaudo et al., 2016). In such systems, multiple adults normally contact offspring, progressively feeding and adjusting nutrition in response to feedback (Field, 2005; Schmickl & Karsai, 2017), making the responses of individual larvae difficult both to follow and interpret.…”

Social Stomach

“…1). Recently, comparable common stomach regulation systems that act in a similar way as the one described above in wasps, have been also identified in honey bees and ants [14][15][16].…”