Memory In Bees Research Articles

Landmark knowledge overrides optic flow in honeybee waggle dance distance estimation.

Honeybees encode in their waggle dances the vector (distance and direction) of an outbound flight to a food source or a new nest site. Optic flow has been identified as the major source of information in the distance estimation. Additional components of distance estimation were also identified, e.g. the sequence of experienced landmarks. Here we address the question of whether bees also use the landscape memory developed during exploratory orientation flights to estimate distance. We took advantage of the fact that flights in a narrow tunnel lead to further distance measures due to higher optic flow. We find that this effect is lost when bees had explored the area in which the tunnel is located and when they have somewhat restricted visual access to the surrounding environment through the mesh on top of the tunnel. These data are interpreted in the context of other findings about the structure of navigational memory in bees that develops during exploratory orientation flights. In particular, the data suggest that bees embed distance measures into a representation of navigational space that stores previously experienced landscape features.

Intake of imidacloprid in lethal and sublethal doses alters gene expression in Apis mellifera bees

Bees are important pollinators for ecosystems and agriculture; however, populations have suffered a decline that may be associated with several factors, including habitat loss, climate change, increased vulnerability to diseases and parasites and use of pesticides. The extensive use of neonicotinoids, including imidacloprid, as agricultural pesticides, leads to their persistence in the environment and accumulation in bees, pollen, nectar, and honey, thereby inducing deleterious effects. Forager honey bees face significant exposure to pesticide residues while searching for resources outside the hive, particularly systemic pesticides like imidacloprid. In this study, 360 Apis mellifera bees, twenty-one days old (supposed to be in the forager phase) previously marked were fed syrup (honey and water, 1:1 m/v) containing a lethal dose (0.081 μg/bee) or sublethal dose (0.00081 μg/bee) of imidacloprid. The syrup was provided in plastic troughs, with 250 μL added per trough onto each plastic Petri dish containing 5 bees (50 μL per bee). The bees were kept in the plastic Petri dishes inside an incubator, and after 1 and 4 h of ingestion, the bees were euthanised and stored in an ultra-freezer (−80 °C) for transcriptome analysis. Following the 1-h ingestion of imidacloprid, 1516 genes (73 from lethal dose; 1509 from sublethal dose) showed differential expression compared to the control, while after 4 h, 758 genes (733 from lethal dose; 25 from sublethal) exhibited differential expression compared to the control. All differentially expressed genes found in the brain tissue transcripts of forager bees were categorised based on gene ontology into functional groups encompassing biological processes, molecular functions, and cellular components. These analyses revealed that sublethal doses might be capable of altering more genes than lethal doses, potentially associated with a phenomenon known as insecticide-induced hormesis. Alterations in genes related to areas such as the immune system, nutritional metabolism, detoxification system, circadian rhythm, odour detection, foraging activity, and memory in bees were present after exposure to the pesticide. These findings underscore the detrimental effects of both lethal and sublethal doses of imidacloprid, thereby providing valuable insights for establishing public policies regarding the use of neonicotinoids, which are directly implicated in the compromised health of Apis mellifera bees.

Microplastics reach the brain and interfere with honey bee cognition

Scientific research on the impact of microplastics (MPs) in terrestrial systems is still emerging, but it has confirmed adverse health effects in organisms exposed to plastics. Although recent studies have shown the toxicological effects of individual MPs polymers on honey bees, the effects of different polymer combinations on cognitive and behavioural performance remain unknown. To fill this knowledge gap, we investigated the effects of oral exposure to spherical MPs on cognitive performance and brain accumulation in the honey bee Apis mellifera. We evaluated the acute toxicity, after a two-day exposure, of polystyrene (PS - 4.8-5.8 μm) and plexiglass (Poly(methyl methacrylate), or PMMA - 1-40 μm) MPs, and a combination of the two (MIX), at two environmentally relevant and one higher concentration (0.5, 5 and 50 mg L−1) and analysed their effects on sucrose responsiveness and appetitive olfactory learning and memory. We also used fluorescent thermoset amino formaldehyde MPs (1–5 μm) to explore whether microspheres of this diameter could penetrate the insect blood-brain barrier (BBB), using Two-Photon Fluorescence Microscopy (TPFM) in combination with an optimized version of the DISCO clearing technique. The results showed that PS reduced sucrose responsiveness, while PMMA had no significant effect; however, the combination had a marked negative effect on sucrose responsiveness. PMMA, PS, and MIX impaired bee learning and memory in bees, with PS showing the most severe effects. 3D brain imaging analysis using TFPM showed that 1–5 μm MPs penetrated and accumulated in the brain after only three days of oral exposure. These results raise concerns about the potential mechanical, cellular, and biochemical damage that MPs may cause to the central nervous system.

Acaricidal Toxicity of Four Essential Oils, Their Predominant Constituents, Their Mixtures against Varroa Mite, and Their Selectivity to Honey Bees (Apis cerana and A. mellifera).

The honey bee (Apis mellifera) faces a significant threat from Varroa destructor, causing the losses of millions of colonies worldwide. While synthetic acaricides are widely used to control Varroa infestations, excessive application has led to resistant strains and poses side effects on the host. Consequently, there is an urgent need for a new acaricide that is both effective and affordable, yet safe to use on bees. One potential source of these acaricides is essential oils (EOs) and their constituents. This study evaluated the acaricidal properties of four essential oils (Eucalyptus globulus, Rosemary officinalis, Trachyspermum ammi (Ethiopian and Indian varieties), their constituents and mixture of constituents against V. destructor through the complete exposure method. Our finding showed that a 1:1 mixture of thymol and carvacrol (4 h-LC50 = 42 μg/mL), thymol (4 h-LC50 = 71 μg/mL), and T. ammi oil (4 h-LC50 = 81-98 μg/mL) were the most toxic test samples against V. destructor. Honey bee behavior and selectivity were also assessed with one additional EO Thymus schimperi, indicating that T. schimperi, T. ammi, and their components were selective and did not affect the learning and memory of bees. In conclusion, the thymol and carvacrol (1:1) mixture was shown to be a promising replacement for synthetic acaricides, being three times more toxic than a commercial acaricide, fluvalinate (4 h-LC50 = 143 μg/mL).

Pollinator cognition: Framing bee memories in an ecological context

Bee memory has been well characterized in laboratory experiments, but its relevance for foraging in an ecological context is less well studied. A new study shows that short-term memory in bumble bees correlates with springtime foraging efficiency, when floral resources are abundant, but not with summer foraging efficiency, when resources are scarce.

Dynamically expressed single ELAV/Hu orthologue elavl2 of bees is required for learning and memory

Changes in gene expression are a hallmark of learning and memory consolidation. Little is known about how alternative mRNA processing, particularly abundant in neuron-specific genes, contributes to these processes. Prototype RNA binding proteins of the neuronally expressed ELAV/Hu family are candidates for roles in learning and memory, but their capacity to cross-regulate and take over each other’s functions complicate substantiation of such links. Honey bees Apis mellifera have only one elav/Hu family gene elavl2, that has functionally diversified by increasing alternative splicing including an evolutionary conserved microexon. RNAi knockdown demonstrates that ELAVL2 is required for learning and memory in bees. ELAVL2 is dynamically expressed with altered alternative splicing and subcellular localization in mushroom bodies, but not in other brain regions. Expression and alternative splicing of elavl2 change during memory consolidation illustrating an alternative mRNA processing program as part of a local gene expression response underlying memory consolidation.

Real-Time Monitoring of DWV-Infected Bee Foraging Behavior Following Histone Deacetylase Inhibitor Treatment

Impairment in learning and memory behaviors in bees due to pesticide exposures and pathogen infections has resulted in massive disappearance of bee populations and subsequent agricultural economic losses. One of the causes of this bee colony collapse disorder (CCD) is the infection of deformed wing virus (DWV). The present study found that sodium butyrate, a histone deacetylase inhibitor (HDACi) previously shown to reverse the detrimental effects of a nicotine-like pesticide on learning behavior and memory in bees, significantly increased the survival of bee after DWV infection and reversed the learning/memory impairment in proboscis extension response (PER) assay. A next-generation sequencing analysis revealed the gene expression profile underwent DWV infection and sodium butyrate treatment and showed that sodium butyrate affected the expression of a wide range of genes involved in diverse biological processes. Particularly, sodium butyrate could up-regulate the expression of genes involved in glycolytic process and memory formation that were down-regulated during DWV infection. As brain physiological analyses presented that sodium butyrate restored the energy supply and the production of neurotransmitter in the infected bees, a large-scale movement tracking was performed by using a wireless sensor network (WSN)-based automatic real-time monitoring system and confirmed that sodium butyrate can improve the ability of DWV-infected bees to return to their hives. In short, we demonstrated the mechanism of how epigenetic regulation can resume the memory function of honeybees and suggested strategies for applying sodium butyrate as possible solutions for reducing the incidence of CCD.

Glyphosate and aminomethylphosphonic acid (AMPA) residues in Brazilian honey

ABSTRACT Glyphosate (GLY) is the most widely used herbicide in the world and studies have shown that its exposure at sublethal doses has led to reduced sensitivity and decreased associative memory in bees. Thus, a high-performance liquid chromatography-fluorescence detector with derivatisation reaction was used to determine residues of GLY and aminomethylphosphonic acid (AMPA) in Brazilian honey. The method met the validation criteria of the European Union (EU) SANTE 11813/2017 guideline and showed a limit of quantitation of 0.04 µg g−1 for both GLY and AMPA. Residues of these compounds were quantitated in honey samples from five Brazilian States. Six samples showed GLY levels above the EU maximum residue limit (0.05 µg g−1) and one sample showed AMPA at 0.10 µg g−1. This study indicates the presence of GLY residues in honey from regions that have had high losses of bee colonies and at the same time, frequent use of GLY in agriculture.

Floral tea polyphenols can improve honey bee memory retention and olfactory sensitivity

Animal-pollinated plants face a common problem, how their defensive anti-herbivore compounds may impair or alter pollinator behavior. Evolution has tailored multiple solutions, which largely involve pollinator tolerance or manipulation, to the benefit of the plant, not the removal of these compounds from pollen or nectar. The tea plant, Camilla sinensis, is famous for the caffeine and tea polyphenols (TP) that it produces in its leaves. However, these compounds are also found in its nectar, which honey bees readily collect. We examined the effects of these compounds on bee foraging choices, learning, memory, and olfactory sensitivity. Foragers preferred a sucrose feeder with 100 µg or 10 µg TP/ml over a control feeder. Caffeine, but not TP, weakly increased honey bee learning. Both caffeine and TP significantly increased memory retention, even when tested 7 d after the last learning trial. In addition, TP generally elevated EAG responsiveness to alarm pheromone odors. These results demonstrate that other secondary plant compounds, not only caffeine, can attract pollinators and influence their learning and memory.

Molecular Recognition of Neonicotinoid Insecticides by Honeybee Nicotinic Receptors and ACHBP Homologues

Neonicotinoids, sometimes called ‘neonics’, are a class of insecticides targeting neuronal nicotinic acetylcholine receptors, which belong to the family of pentameric ligand-gated ion channels (pLGICs) or Cys-loop receptors. The widespread application of these neurotoxic insecticides in agriculture is tied to the worldwide decline of bee populations, due to their sublethal effects on bee memory, behavior and reproduction. In 2018, the member states of the European Union, agreed on a ban of most neonicotinoids, with exemption for greenhouses. However, a new generation of sulfoximine-based insecticides, such as sulfoxaflor, has recently been introduced to the agricultural market claiming they are chemically distinct from neonicotinoids. Using the acetylcholine binding protein (AChBP) as a well-established tool for structural studies of nicotinic receptors, we engineered AChBP variants that mimic the neurotransmitter binding site of honeybee nicotinic receptors. In parallel, we pursued functional characterization of these receptors expressed in Xenopus oocytes and using electrophysiological techniques. Three-dimensional structures of sulfoxaflor-bound AChBPs reveal that the molecule is recognized through receptor interactions that are virtually identical to the neonicotinoid thiacloprid. Surprisingly, binding assays reveal a ∼100-fold difference in affinity between sulfoxaflor and thiacloprid, leaving open important questions with respect to the precise molecular mechanism of sulfoxaflor recognition. Our study will attempt to begin addressing these central questions.

Inhibitory learning of phototaxis by honeybees in a passive-avoidance task.

Honeybees are a standard model for the study of appetitive learning and memory. Yet, fewer attempts have been performed to characterize aversive learning and memory in this insect and uncover its molecular underpinnings. Here, we took advantage of the positive phototactic behavior of bees kept away from the hive in a dark environment and established a passive-avoidance task in which they had to suppress positive phototaxis. Bees placed in a two-compartment box learned to inhibit spontaneous attraction to a compartment illuminated with blue light by associating and entering into that chamber with shock delivery. Inhibitory learning resulted in an avoidance memory that could be retrieved 24 h after training and that was specific to the punished blue light. The memory was mainly operant but involved a Pavlovian component linking the blue light and the shock. Coupling conditioning with transcriptional analyses in key areas of the brain showed that inhibitory learning of phototaxis leads to an up-regulation of the dopaminergic receptor gene Amdop1 in the calyces of the mushroom bodies, consistently with the role of dopamine signaling in different forms of aversive learning in insects. Our results thus introduce new perspectives for uncovering further cellular and molecular underpinnings of aversive learning and memory in bees. Overall, they represent an important step toward comparative learning studies between the appetitive and the aversive frameworks.

The Molecular Recognition Of The Insecticide Sulfoxaflor By Nicotinic Receptors Is Based On Atomic Interactions Identical To Bee‐Harming Neonicotinoids

Neonicotinoids, sometimes called ‘neonics’, are a class of insecticides targeting neuronal nicotinic acetylcholine receptors, which belong to the family of pentameric ligand-gated ion channels (pLGICs) or Cys-loop receptors. The widespread application of these neurotoxic insecticides in agriculture is tied to the worldwide decline of bee populations, due to their sublethal effects on bee memory, behavior and reproduction. In 2018, the member states of the European Union, agreed on a ban of most neonicotinoids, with exemption for greenhouses. However, a new generation of sulfoximine-based insecticides, such as sulfoxaflor, has recently been introduced to the agricultural market claiming they are chemically distinct from neonicotinoids. Using the acetylcholine binding protein (AChBP) as a well-established tool for structural studies of nicotinic receptors, we engineered AChBP variants that reliably mimic the neurotransmitter binding site of honeybee nicotinic receptors. Three-dimensional structures of sulfoxaflor-bound receptors reveal, despite the distinct chemistry claim, that the molecule is recognized through receptor interactions that are virtually identical to neonicotinoids. This observation suggests that sulfoximine- and neonicotinoid-type insecticides share a common mode of action, and that the legislative ban on certain insecticides should, as a precaution, be expanded based on pharmacological activity at their target receptors rather than on chemical composition. Crystal structure of an insect nicotinic receptor mimic (iAChBP) in complex with the insecticide sulfoxaflor. This abstract is from the Experimental Biology 2019 Meeting. There is no full text article associated with this abstract published in The FASEB Journal.

Length of stimulus presentation and visual angle are critical for efficient visual PER conditioning in the restrained honey bee, Apis mellifera.

Learning visual cues is an essential capability of bees for vital behaviors such as orientation in space and recognition of nest sites, food sources and mating partners. To study learning and memory in bees under controlled conditions, the proboscis extension response (PER) provides a well-established behavioral paradigm. While many studies have used the PER paradigm to test olfactory learning in bees because of its robustness and reproducibility, studies on PER conditioning of visual stimuli are rare. In this study, we designed a new setup to test the learning performance of restrained honey bees and the impact of several parameters: stimulus presentation length, stimulus size (i.e. visual angle) and ambient illumination. Intact honey bee workers could successfully discriminate between two monochromatic lights when the color stimulus was presented for 4, 7 and 10 s before a sugar reward was offered, reaching similar performance levels to those for olfactory conditioning. However, bees did not learn at shorter presentation durations. Similar to free-flying honey bees, harnessed bees were able to associate a visual stimulus with a reward at small visual angles (5deg) but failed to utilize the chromatic information to discriminate the learned stimulus from a novel color. Finally, ambient light had no effect on acquisition performance. We discuss possible reasons for the distinct differences between olfactory and visual PER conditioning.

Quantifying the impact of pesticides on learning and memory in bees.

Most insecticides are insect neurotoxins. Evidence is emerging that sublethal doses of these neurotoxins are affecting the learning and memory of both wild and managed bee colonies, exacerbating the negative effects of pesticide exposure and reducing individual foraging efficiency.Variation in methodologies and interpretation of results across studies has precluded the quantitative evaluation of these impacts that is needed to make recommendations for policy change. It is not clear whether robust effects occur under acute exposure regimes (often argued to be more field‐realistic than the chronic regimes upon which many studies are based), for field‐realistic dosages, and for pesticides other than neonicotinoids.Here we use meta‐analysis to examine the impact of pesticides on bee performance in proboscis extension‐based learning assays, the paradigm most commonly used to assess learning and memory in bees. We draw together 104 (learning) and 167 (memory) estimated effect sizes across a diverse range of studies.We detected significant negative effects of pesticides on learning and memory (i) at field realistic dosages, (ii) under both chronic and acute application, and (iii) for both neonicotinoid and non‐neonicotinoid pesticides groups.We also expose key gaps in the literature that include a critical lack of studies on non‐Apis bees, on larval exposure (potentially one of the major exposure routes), and on performance in alternative learning paradigms. Policy implications. Procedures for the registration of new pesticides within EU member states now typically require assessment of risks to pollinators if potential target crops are attractive to bees. However, our results provide robust quantitative evidence for subtle, sublethal effects, the consequences of which are unlikely to be detected within small‐scale prelicensing laboratory or field trials, but can be critical when pesticides are used at a landscape scale. Our findings highlight the need for long‐term postlicensing environmental safety monitoring as a requirement within licensing policy for plant protection products.

Larval exposure to thiamethoxam and American foulbrood: effects on mortality and cognition in the honey bee Apis mellifera

Here, we examined the in vitro effects of co-exposure to a pathogen and a common neonicotinoid on honey bee larvae survival and on adult learning behavior following a standard olfactory conditioning procedure based on the proboscis extension response paradigm. We exposed or co-exposed honey bee larvae to American foulbrood and to sub-lethal doses of thiamethoxam (chronic exposure). Our results revealed no additive effects between the two stressors on larval mortality. However, the present work provides the first evidence of impaired learning and memory in adult bees that were fed thiamethoxam (0.6 ng/bee) during the larval stage. We also show no alterations in learning and memory in bees after infection with American foulbrood at the larval stage. The present study contributes to our knowledge of the sub-lethal effects of neonicotinoids on honey bee larvae and adults.

Low doses of neonicotinoid pesticides in food rewards impair short-term olfactory memory in foraging-age honeybees.

Neonicotinoids are often applied as systemic seed treatments to crops and have reported negative impact on pollinators when they appear in floral nectar and pollen. Recently, we found that bees in a two-choice assay prefer to consume solutions containing field-relevant doses of the neonicotinoid pesticides, imidacloprid (IMD) and thiamethoxam (TMX), to sucrose alone. This suggests that neonicotinoids enhance the rewarding properties of sucrose and that low, acute doses could improve learning and memory in bees. To test this, we trained foraging-age honeybees to learn to associate floral scent with a reward containing nectar-relevant concentrations of IMD and TMX and tested their short (STM) and long-term (LTM) olfactory memories. Contrary to our predictions, we found that none of the solutions enhanced the rate of olfactory learning and some of them impaired it. In particular, the effect of 10 nM IMD was observed by the second conditioning trial and persisted 24 h later. In most other groups, exposure to IMD and TMX affected STM but not LTM. Our data show that negative impacts of low doses of IMD and TMX do not require long-term exposure and suggest that impacts of neonicotinoids on olfaction are greater than their effects on rewarding memories.

USING AN IMPROVED BEE MEMORY DIFFERENTIAL EVOLUTION ALGORITHM FOR PARAMETER ESTIMATION TO SIMULATE BIOCHEMICAL PATHWAYS

When analyzing a metabolic pathway in a mathematical model, it is important that the essential parameters are estimated correctly. However, this process often faces few problems like when the number of unknown parameters increase, trapping of data in the local minima, repeated exposure to bad results during the search process and occurrence of noisy data. Thus, this paper intends to present an improved bee memory differential evolution (IBMDE) algorithm to solve the mentioned problems. This is a hybrid algorithm that combines the differential evolution (DE) algorithm, the Kalman filter, artificial bee colony (ABC) algorithm, and a memory feature. The aspartate and threonine biosynthesis pathway, and cell cycle pathway are the metabolic pathways used in this paper. For three production simulation pathways, the IBMDE managed to robustly produce the estimated optimal kinetic parameter values with significantly reduced errors. Besides, it also demonstrated faster convergence time compared to the Nelder–Mead (NM), simulated annealing (SA), the genetic algorithm (GA) and DE, respectively. Most importantly, the kinetic parameters that were generated by the IBMDE have improved the production rates of desired metabolites better than other estimation algorithms. Meanwhile, the results proved that the IBMDE is a reliable estimation algorithm.

Parameter Estimation by Using an Improved Bee Memory Differential Evolution Algorithm (IBMDE) to Simulate Biochemical Pathways

Assessing and estimating essential parameters for a metabolic pathway by using a mathematical model is a significant step in Systems Biology. However, estimating process often faces numerous obstacles, for example when the number of unknown parameters escalates or data has noise, gets trapped in local minima and or having repeated exploration of poor solution during search process. Thus, this study proposes an improved Bee Memory Differential Evolution algorithm (IBMDE), which is a combination of the Differential Evolution algorithm (DE), the Kalman Filter (KF), the Artificial Bee Colony algorithm (ABC), and a memory feature to solve the aforementioned problems. The implemented metabolic pathways for this improved estimation algorithm were glycerol and pyruvate synthesis pathways. IBMDE was successful in generating the estimated optimal kinetic parameter values with noticeable reduction in errors (81.36% and 99.46% respectively) and faster convergence times (6.19% and 15.72% respectively) compared to DE, the Genetic Algorithm (GA), the Nelder Mead (NM), and the Simulated Annealing (SA). The results indicated that, most importantly, the kinetic parameters produced by IBMDE had enhanced the production of desired metabolites than the other estimation algorithms. Besides that, the results also demonstrated the reliability of IBMDE as an estimation algorithm in terms of lower error.

Caffeine Boosts Bees' Memories

Caffeine in floral nectar enhances the memory of bees for the flowers' scent by altering response properties of neurons in the bee brain. [Also see Report by Wright et al. ]

Restoring the males’ honor: Pavlovian PER conditioning of visual cues in bumblebee drones

Event Abstract Back to Event Restoring the males’ honor: Pavlovian PER conditioning of visual cues in bumblebee drones Leonie Lichtenstein1, Frank M. Sommerlandt1* and Johannes Spaethe1 1 University of Würzburg, Department of Behavioral Physiology and Sociobiology, Germany Learning visual cues is essential in bees for orientation, as well as for the recognition of nest sites, food sources and mating partners. A well-established method for a quantitative evaluation of learning and memory in bees under controlled environmental conditions is the proboscis extension response (PER). After several decades of failure, the PER assay has recently been successfully applied for the associative conditioning of visual stimuli in intact honeybees (Dobrin and Fahrbach, 2012) and bumblebees (Riveros and Gronenberg, 2012). Research using the PER method has primarily focused on workers, whereas males were largely neglected. In the present study, we tested intact Bombus terrestris drones in different visual learning tasks using classical PER conditioning and different monochromatic light stimuli (435, 488, and 528nm). Individuals were confronted with absolute (A+) and differential (A+ vs. B-) conditioning tasks. Furthermore, drones were tested to determine if they can retrieve the learned stimulus after two hours and transfer this information to a subsequent discrimination task in a Y-maze. Our results indicate that drones are capable of discriminating between monochromatic light stimuli, and they attain performance levels similar to that of workers. This approach will allow us to investigate color discrimination abilities of bees in different parts of the color space by increasing the color similarity between stimuli in differential conditioning under controlled conditions. Comparing the perceptual capabilities of different bee species using the PER paradigm may thus enable us to better understand the evolution of color processing within the family Apidae. Figure 1: Probability of proboscis extension response (PER) during absolute conditioning of three color stimuli. Bumblebee drones were trained to associate a color stimulus of particular wavelength (435nm, n=20; 488nm, n=21; and 528nm, n=17) with a sucrose reward. After two hours, a transfer test showed that drones can retrieve the learned color (conditioned stimulus; CS) and discriminate against a novel color (Ncol). Learning performance did not differ among colors. Figure 1 Keywords: Bumblebee, associative learning, proboscis extension response, monochromatic light, visual conditioning, Bombus terrestris, memory consolidation, Drones Conference: International Conference on Invertebrate Vision, Fjälkinge, Sweden, 1 Aug - 8 Aug, 2013. Presentation Type: Poster presentation preferred Topic: Colour and polarisation vision Citation: Lichtenstein L, Sommerlandt FM and Spaethe J (2019). Restoring the males’ honor: Pavlovian PER conditioning of visual cues in bumblebee drones. Front. Physiol. Conference Abstract: International Conference on Invertebrate Vision. doi: 10.3389/conf.fphys.2013.25.00107 Copyright: The abstracts in this collection have not been subject to any Frontiers peer review or checks, and are not endorsed by Frontiers. They are made available through the Frontiers publishing platform as a service to conference organizers and presenters. The copyright in the individual abstracts is owned by the author of each abstract or his/her employer unless otherwise stated. Each abstract, as well as the collection of abstracts, are published under a Creative Commons CC-BY 4.0 (attribution) licence (https://creativecommons.org/licenses/by/4.0/) and may thus be reproduced, translated, adapted and be the subject of derivative works provided the authors and Frontiers are attributed. For Frontiers’ terms and conditions please see https://www.frontiersin.org/legal/terms-and-conditions. Received: 28 Feb 2013; Published Online: 09 Dec 2019. * Correspondence: Mr. Frank M Sommerlandt, University of Würzburg, Department of Behavioral Physiology and Sociobiology, Würzburg, 97074, Germany, frank.sommerlandt@uni-wuerzburg.de Login Required This action requires you to be registered with Frontiers and logged in. To register or login click here. Abstract Info Abstract The Authors in Frontiers Leonie Lichtenstein Frank M Sommerlandt Johannes Spaethe Google Leonie Lichtenstein Frank M Sommerlandt Johannes Spaethe Google Scholar Leonie Lichtenstein Frank M Sommerlandt Johannes Spaethe PubMed Leonie Lichtenstein Frank M Sommerlandt Johannes Spaethe Related Article in Frontiers Google Scholar PubMed Abstract Close Back to top Javascript is disabled. Please enable Javascript in your browser settings in order to see all the content on this page.