The honeybee as a model for understanding the basis of cognition

Key Points

  • Honeybees with their tiny brains exhibit complex social and navigational behaviours and possess a relatively rich cognitive repertoire. A unique feature of honeybee behaviour is the waggle dance, a ritualized movement that communicates locations and their properties.

  • The honeybee brain consists of 1 million neurons that are structured in highly ordered neuropils. Many of the central neurons are individually identifiable and some of them are registered in a three-dimensional virtual standard brain atlas.

  • Cognitive forms of learning in the bee include categorization, extraction of dependences on context, sequences and combinations, and evaluation of sequential reward values.

  • The search for neural correlates of learning and memory processing in the honeybee is facilitated by a highly versatile behavioural paradigm: the classical conditioning of the proboscis extension response. This paradigm allows the monitoring of neural events in defined neural networks and single neurons together with behavioural change.

  • Learning-related plasticity is found in all neural components of the olfactory pathway and in an identified reward neuron. The sparse and combinatorial code of odours is predominantly enhanced for the learned odour at the input site of the mushroom body, whereas the mushroom body output codes the value of the learned signals.

  • Memory as characterized by behavioural and molecular studies is processed in four distinct phases in honeybees, which is similar to common properties of memory in other animal species.


Honeybees contradict the notion that insect behaviour tends to be relatively inflexible and stereotypical. Indeed, they live in colonies and exhibit complex social, navigational and communication behaviours, as well as a relatively rich cognitive repertoire. Because these relatively complex behaviours are controlled by a brain consisting of only 1 million or so neurons, honeybees offer an opportunity to study the relationship between behaviour and cognition in neural networks that are limited in size and complexity. Most recently, the honeybee has been used to model learning and memory formation, highlighting its utility for neuroscience research, in particular for understanding the basis of cognition.

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Figure 1: Projection pattern of VUMmx1, which encodes reward in olfactory learning.
Figure 2: Memory formation is a multi-phase process in honeybees.
Figure 3: Visualizing the engram of learned odours at the input side of the mushroom body.
Figure 4: Brain structures and their connections housing the olfactory memory trace.


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I am grateful to D. Eisenhardt, B. Brembs, J. Rybak and G. Leboulle for commenting on an earlier version of the manuscript. I am particularly grateful to G. Leboulle for advice on the molecular genetic studies in honeybees, and to J. Rybak for his comments about the anatomy of the insect brain. My work is supported by the Deutsche Forschungsgemeinschaft, Gemeinnützige Stiftung Hertie and Klaus Tschira Stiftung.

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Menzel, R. The honeybee as a model for understanding the basis of cognition. Nat Rev Neurosci 13, 758–768 (2012).

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