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. 2019 Jul 16;9:10313. doi: 10.1038/s41598-019-46482-4

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© The Author(s) 2019

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons license, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/.

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Responsiveness and learning protocols for appetitive and aversive hedonic modalities performed on the same individuals (n = 226). (A) Heat responsiveness. Red circles, %SER to a series of increasing temperatures; white circles, %SER of the same bees to the presentation of a tactile (unheated) stimulus (control). (B) Sucrose responsiveness. Blue circles, %PER to a series of sucrose solutions of increasing concentration; white circles, %PER of the same bees to the presentation of water (control). (C) Differential aversive conditioning of the SER. Red circles, %SER to the CS+ (reinforced odor) along the 8 trials; white circles, %SER to the CS− (non-reinforced odor). (D) Differential appetitive conditioning of the PER. Blue circles, %PER to the CS+ along the 8 trials; white circles, %PER to the CS−. (ANOVA for repeated measurements, AB; stimulus x concentration or CD: stimulus x trial interaction, ***p < 0.001).