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Biological patterns are often constructed via a combination of mechanisms including self-organization, templates and recipes. Our understanding of self-organization is becoming increasingly clear, yet how multiple mechanisms work together and what selective advantage they confer over simpler mechanisms is poorly understood. Honeybee (Apis mellifera) combs exhibit a pattern of brood at the bottom, pollen in a band next to it and honey at the top. This study constructs an agent-based model, derived from experimental studies, to determine both how self-organization interacts with two templates and to elucidate a selective basis for the use of multiple mechanisms. The vertical pattern of honey and brood is shown to be dependent on a gravity-based template, while the pollen band is shown to form via the interaction of a queen-based template and self-organization. The study suggests that the selective basis for this complex mechanism may be that colonies have higher growth rates when multiple mechanisms are used as opposed to self-organization alone. As self-organization is used in many contexts in which the addition of supplemental mechanisms could be advantageous, this result may be of general significance to many biological systems.

Pattern formation on the combs of honeybees: increasing fitness by coupling self-organization with templates