Search

Bees are flexible and adaptive learners, capable of learning stimuli seen on arrival and at departure from flowers where they have fed. This gives bees the potential to learn all information associated with a feeding event, but it also presents the challenge of managing information that is irrelevant, inconsistent, or conflicting. Here, we examined how presenting bumblebees with conflicting visual information before and after feeding influenced their learning rate and what they learned. Bees were trained to feeder stations mounted in front of a computer monitor. Visual stimuli were displayed behind each feeder station on the monitor. Positively reinforced stimuli (CS +) marked feeders offering sucrose solution. Negatively reinforced stimuli (CS-) marked feeders offering quinine solution. While alighted at the feeder station the stimuli were likely not visible to the bee. The "constant stimulus" training group saw the same stimulus throughout. For the "switched stimulus" training group, the CS + changed to the CS- during feeding. Learning was slower in the "switched stimulus" training group compared to the constant stimulus" group, but the training groups did not differ in their learning performance or the extent to which they generalised their learning. The information conflict in the "switched stimulus" group did not interfere with what had been learned. Differences between the "switched" and "constant stimulus" groups were greater for bees trained on a horizontal CS + than a vertical CS + suggesting bees differ in their processing of vertically and horizontally oriented stimuli. We discuss how bumblebees might resolve this type of information conflict so effectively, drawing on the known neurobiology of their visual learning system., Competing Interests: Declarations. Conflict of interest: The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. Ethical approval: Our research involved bumblebees from commercially available colonies dedicated to research for which an approval of an ethical committee is not mandatory. The protocols comply with standard welfare practice in our field and a minimum number of individuals were used to study our scientific question. The animals were not harmed during the experimental procedures., (© 2025. The Author(s).)

Background: Recent articles have described the challenges in developing nurse leaders to advance nursing science, improve health, healthcare, and health equity, and build the next generation of nurses. Over the past 25 years, the Robert Wood Johnson Foundation (RWJF) has implemented many programs to develop nurse leaders to address these challenges., Purpose: This article shares lessons learned from four RWJF programs and how the authors recommend those lessons can be applied today., Methods: The Robert Wood Johnson Foundation (RWJF) has supported nursing leadership development for over fifty years. Discussion of four RWJF nursing programs including the Executive Nurse Fellows (1998-2017); the New Jersey Nursing Initiative (2009-2016); the Nurse Faculty Scholars (2007-2017) and the Future of Nursing Scholars (2013-2023) is used to highlight strategies and lessons learned., Findings: Key lessons for developing the next generation of nurse leaders and strengthening the nurse leader pipeline include the importance of continuing leadership development over the course of one's career; accelerating the nursing PhD to faculty pipeline; strong intentional mentoring; and building supportive communities to ensure long term impact., Discussion: While these program examples had purposeful philanthropic funding, the lessons can also be implemented by building partnerships with federal agencies, philanthropy, and universities., Competing Interests: Declaration of Competing Interest The authors declare no conflicts of interest., (Copyright © 2024 The Author(s). Published by Elsevier Inc. All rights reserved.)

The evolution of cognition can be understood in terms of a few major transitions-changes in the computational architecture of nervous systems that changed what cognitive capacities could be evolved by downstream lineages. We demonstrate how the idea of a major cognitive transition can be modeled in terms of where a system's effective computational architecture falls on the well-studied hierarchy of formal automata (HFA). We then use recent work connecting artificial neural networks to the HFA, which provides a way to make the structure-architecture link in natural systems. We conclude with reflections on the power and the challenges of traditional thinking when applied to neural architectures. This article is categorized under: Cognitive Biology > Evolutionary Roots of Cognition Psychology > Comparative Philosophy > Foundations of Cognitive Science., (© 2024 The Authors. WIREs Cognitive Science published by Wiley Periodicals LLC.)

The thriving field of comparative cognition examines the behaviour of diverse animals in cognitive terms. Comparative cognition research has primarily focused on the abilities of animals - what tasks they can do - rather than on the limits of their cognition - tasks that exceed an animal's cognitive abilities. We propose that understanding and identifying cognitive limits is as important as demonstrating the capacities of animal minds. Here, we identify challenges that have deterred the study of cognitive limits related to epistemic, practical and publication problems. The epistemic problem is concerned with how we can confidently infer a cognitive limit from null or negative results. The practical problem is how can we be certain our research has identified a cognitive limit rather than failures in tasks due to methodological or experimental design issues. The publication problem outlines the publication bias toward positive and exciting results over negative or null results in animal cognition. We propose solutions to these three challenges and examples of how to conduct research to confidently identify and confirm cognitive limits in animals. We believe a refocus on the cognitive limits of animals is the next step in the field of comparative cognition. Knowing the limits to the intelligence of different animals will aid us in appreciating the diversity of animal intelligence, and will resolve outstanding questions of how cognition evolves., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2024 Elsevier Inc. All rights reserved.)

Laboratory studies show detrimental effects of metallic pollutants on invertebrate behaviour and cognition, even at low levels. Here we report a field study on Western honey bees exposed to metal and metalloid pollution through dusts, food and water at a historic mining site. We analysed more than 1000 bees from five apiaries along a gradient of contamination within 11 km of a former gold mine in Southern France. Bees collected close to the mine exhibited olfactory learning performances lower by 36% and heads smaller by 4%. Three-dimensional scans of bee brains showed that the olfactory centres of insects sampled close to the mine were also 4% smaller, indicating neurodevelopmental issues. Our study raises serious concerns about the health of honey bee populations in areas polluted with potentially harmful elements, particularly with arsenic, and illustrates how standard cognitive tests can be used for risk assessment., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2023 The Authors. Published by Elsevier B.V. All rights reserved.)

The comparative approach is a powerful way to explore the relationship between brain structure and cognitive function. Thus far, the field has been dominated by the assumption that a bigger brain somehow means better cognition. Correlations between differences in brain size or neuron number between species and differences in specific cognitive abilities exist, but these correlations are very noisy. Extreme differences exist between clades in the relationship between either brain size or neuron number and specific cognitive abilities. This means that correlations become weaker, not stronger, as the taxonomic diversity of sampled groups increases. Cognition is the outcome of neural networks. Here we propose that considering plausible neural network models will advance our understanding of the complex relationships between neuron number and different aspects of cognition. Computational modelling of networks suggests that adding pathways, or layers, or changing patterns of connectivity in a network can all have different specific consequences for cognition. Consequently, models of computational architecture can help us hypothesise how and why differences in neuron number might be related to differences in cognition. As methods in connectomics continue to improve and more structural information on animal brains becomes available, we are learning more about natural network structures in brains, and we can develop more biologically plausible models of cognitive architecture. Natural animal diversity then becomes a powerful resource to both test the assumptions of these models and explore hypotheses for how neural network structure and network size might delimit cognitive function., (© 2023 S. Karger AG, Basel.)

The evolutionary history of animal cognition appears to involve a few major transitions: major changes that opened up new phylogenetic possibilities for cognition. Here, we review and contrast current transitional accounts of cognitive evolution. We discuss how an important feature of an evolutionary transition should be that it changes what is evolvable, so that the possible phenotypic spaces before and after a transition are different. We develop an account of cognitive evolution that focuses on how selection might act on the computational architecture of nervous systems. Selection for operational efficiency or robustness can drive changes in computational architecture that then make new types of cognition evolvable. We propose five major transitions in the evolution of animal nervous systems. Each of these gave rise to a different type of computational architecture that changed the evolvability of a lineage and allowed the evolution of new cognitive capacities. Transitional accounts have value in that they allow a big-picture perspective of macroevolution by focusing on changes that have had major consequences. For cognitive evolution, however, we argue it is most useful to focus on evolutionary changes to the nervous system that changed what is evolvable, rather than to focus on specific cognitive capacities.

Honey bee ecology demands they make both rapid and accurate assessments of which flowers are most likely to offer them nectar or pollen. To understand the mechanisms of honey bee decision-making, we examined their speed and accuracy of both flower acceptance and rejection decisions. We used a controlled flight arena that varied both the likelihood of a stimulus offering reward and punishment and the quality of evidence for stimuli. We found that the sophistication of honey bee decision-making rivalled that reported for primates. Their decisions were sensitive to both the quality and reliability of evidence. Acceptance responses had higher accuracy than rejection responses and were more sensitive to changes in available evidence and reward likelihood. Fast acceptances were more likely to be correct than slower acceptances; a phenomenon also seen in primates and indicative that the evidence threshold for a decision changes dynamically with sampling time. To investigate the minimally sufficient circuitry required for these decision-making capacities, we developed a novel model of decision-making. Our model can be mapped to known pathways in the insect brain and is neurobiologically plausible. Our model proposes a system for robust autonomous decision-making with potential application in robotics., Competing Interests: HM, JM, ND, AB No competing interests declared, (© 2023, MaBouDi et al.)

The salivary gland of the black field cricket, Teleogryllus commodus Walker changed size between being starved and fed. Crickets without access to food for 72 h showed a reduction in both wet and dry mass of the glands compared with the glands from continuously fed animals at 72 h. Glands returned to size following ingestion within 10 min. Salivary glands of starved crickets (72 h) were incubated in saline containing either serotonin (5-HT) or dopamine (DA). Glands increased to pre-starvation size after 1 h incubation in situ with either 10 -4 moles L -1 5-HT or 10 -4 moles L -1 DA, although lower concentrations (10 -5 moles L -1 ) did not affect gland size. From immunohistochemistry, amines appeared to shift from zymogen cells during starvation to parietal cells following feeding. High-performance liquid chromatography showed that serotonin concentration is higher than dopamine in the salivary gland removed from starved and fed crickets, but the quantity of these compounds was not dependent upon feeding state; the amine quantities increased as gland size increased. Further work is necessary to determine what might be the stimulus for gland growth and if dopamine and serotonin play a role in the stimulation of salivary gland growth after a period of starvation.

The role of the epigenome in phenotypic plasticity is unclear presently. Here we used a multiomics approach to explore the nature of the epigenome in developing honey bee (Apis mellifera) workers and queens. Our data clearly showed distinct queen and worker epigenomic landscapes during the developmental process. Differences in gene expression between workers and queens become more extensive and more layered during the process of development. Genes known to be important for caste differentiation were more likely to be regulated by multiple epigenomic systems than other differentially expressed genes. We confirmed the importance of two candidate genes for caste differentiation by using RNAi to manipulate the expression of two genes that differed in expression between workers and queens were regulated by multiple epigenomic systems. For both genes the RNAi manipulation resulted in a decrease in weight and fewer ovarioles of newly emerged queens compared to controls. Our data show that the distinct epigenomic landscapes of worker and queen bees differentiate during the course of larval development., Competing Interests: Declaration of competing interest The authors declare that they have no competing interests., (Copyright © 2023 Elsevier Ltd. All rights reserved.)

As you are reading this text, your eyes scan across the page, even if you keep your head perfectly still. New research reveals that flies can perform analogous retinal movements, despite their eyes being rigidly fixed to their heads., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2022 Elsevier Inc. All rights reserved.)

Are animals' preferences determined by absolute memories for options (e.g. reward sizes) or by their remembered ranking (better/worse)? The only studies examining this question suggest humans and starlings utilise memories for both absolute and relative information. We show that bumblebees' learned preferences are based only on memories of ordinal comparisons. A series of experiments showed that after learning to discriminate pairs of different flowers by sucrose concentration, bumblebees preferred flowers (in novel pairings) with (1) higher ranking over equal absolute reward, (2) higher ranking over higher absolute reward, and (3) identical qualitative ranking but different quantitative ranking equally. Bumblebees used absolute information in order to rank different flowers. However, additional experiments revealed that, even when ranking information was absent (i.e. bees learned one flower at a time), memories for absolute information were lost or could no longer be retrieved after at most 1 hr. Our results illuminate a divergent mechanism for bees (compared to starlings and humans) of learned preferences that may have arisen from different adaptations to their natural environment., Competing Interests: CS, YZ, YF, YL, MR, LS, RR, LC, AB, FP No competing interests declared, (© 2022, Solvi, Zhou et al.)

Much of human cognition involves two different types of reasoning that operate together. Type 1 reasoning systems are intuitive and fast, whereas Type 2 reasoning systems are reflective and slow. Why has our cognition evolved with these features? Both systems are coherent and in most ecological circumstances either alone is capable of coming up with the right answer most of the time. Neural tissue is costly, and thus far evolutionary models have struggled to identify a benefit of operating two systems of reasoning. To explore this issue we take a broad comparative perspective. We discuss how dual processes of cognition have enabled the emergence of selective attention in insects, transforming the learning capacities of these animals. Modern AIs using dual systems of learning are able to learn how their vast world works and how best to interact with it, allowing them to exceed human levels of performance in strategy games. We propose that the core benefits of dual processes of reasoning are to narrow down a problem space in order to focus cognitive resources most effectively., (Copyright © 2022 The Authors. Published by Elsevier B.V. All rights reserved.)

Whether animals can actively avoid food contaminated with harmful compounds through taste is key to assess their ecotoxicological risks. Here, we investigated the ability of honey bees to perceive and avoid food resources contaminated with common metal pollutants known to impair behaviour at low concentrations. In laboratory assays, bees did not discriminate food contaminated with arsenic, lead or zinc and ingested it readily, up to estimated doses of 929.1 μg g -1 As, 6.45 mg g -1 Pb and 72.46 mg g -1 Zn. A decrease of intake and appetitive responses indicating metal detection was only observed at the highest concentrations of lead (3.6 mM) and zinc (122.3 mM) through contact with the antennae and the proboscis. Electrophysiological analyses confirmed that only high concentrations of the three metals in a sucrose solution induced a consistently reduced neural response to sucrose in antennal taste receptors (As: >0.1 μM, Pb: >1 mM; Zn: >100 mM). Overall, cellular and behavioural responses did not provide evidence for specific mechanisms that would support selective detection of toxic metals (arsenic, lead), as compared to zinc, which has important biological functions. Our results thus show that honey bees can avoid metal pollutants in their food only at high concentrations unlikely to be encountered in the environment. By contrast, they appear to be unable to detect low, yet harmful, concentrations found in flowers. Metal pollution at trace levels is therefore a major threat for pollinators., (Copyright © 2022 Elsevier Ltd. All rights reserved.)

Volatile odors from flowers play an important role in plant-pollinator interaction. The honeybee is an important generalist pollinator of many plants. Here, we explored whether any components of the odors of a range of honeybee-pollinated plants are commonly involved in the interaction between plants and honeybees. We used a needle trap system to collect floral odors, and GC-MS analysis revealed nonanal was the only component scent detected in 12 different honeybee-pollinated flowers and not present in anemophilous plant species. For Ligustrum compactum, blooming flowers released significantly more nonanal than buds and faded flowers. For Sapium sebiferum, nonanal release through the day correlated with nectar secretion. Experimentally increasing nectar load in flowers of Sapium sebiferum, Ligustrum compactum, and Castanea henryi increased nonanal levels also. Nonanal was also detected in flower nectar and honeys from experimental colonies. Electroantennogram recordings and behavioral observations showed that untrained honeybees could detect and were strongly attracted to nonanal. We argue that nonanal persists in both honey and nectar odors facilitating a learned association between nonanal and food reward in honeybees., (© 2022. The Author(s), under exclusive licence to Springer-Verlag GmbH Germany, part of Springer Nature.)

The distinct honeybee ( Apis mellifera ) worker and queen castes have become a model for the study of genomic mechanisms of phenotypic plasticity. Here we performed a nanopore-based direct RNA sequencing with exceptionally long reads to compare the mRNA transcripts between queen and workers at three points during their larval development. We found thousands of significantly differentially expressed transcript isoforms (DEIs) between queen and worker larvae. These DEIs were formatted by a flexible splicing system. We showed that poly(A) tails participated in this caste differentiation by negatively regulating the expression of DEIs. Hundreds of isoforms uniquely expressed in either queens or workers during their larval development, and isoforms were expressed at different points in queen and worker larval development demonstrating a dynamic relationship between isoform expression and developmental mechanisms. These findings show the full complexity of RNA processing and transcript expression in honey bee phenotypic plasticity., Competing Interests: The authors declare no competing interest., (© 2022 The Authors.)

Understanding the cumulative risk of chemical mixtures at environmentally realistic concentrations is a key challenge in honey bee ecotoxicology. Ecotoxicogenomics, including transcriptomics, measures responses in individual organisms at the molecular level which can provide insights into the mechanisms underlying phenotypic responses induced by one or more stressors and link impacts on individuals to populations. Here, fifth instar honey bee larvae were sampled from a previously reported field experiment exploring the phenotypic impacts of environmentally realistic chronic exposures of the pesticide imidacloprid (5 μg.kg -1 for six weeks) and the acaricide thymol (250 g.kg -1 applied via Apiguard gel in-hive for four weeks), both separately and in combination. RNA-seq was used to discover individual and interactive chemical effects on larval gene expression and to uncover molecular mechanisms linked to reported adult and colony phenotypes. The separate and combined treatments had distinct gene expression profiles which represented differentially affected signaling and metabolic pathways. The molecular signature of the mixture was characterised by additive interactions in canonical stress responses associated with oxidative stress and detoxification, and non-additive interactions in secondary responses including developmental, neurological, and immune pathways. Novel emergent impacts on eye development genes correlated with long-term defects in visual learning performance as adults. This is consistent with these chemicals working through independent modes of action that combine to impact common downstream pathways, and highlights the importance of establishing mechanistic links between molecular and phenotypic responses when predicting effects of chemical mixtures on ecologically relevant population outcomes., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2022 Commonwealth Scientific and Industrial Research Organisation. Published by Elsevier B.V. All rights reserved.)

How neonicotinoid contamination affects honey bees remains controversial. Studies have yielded contradictory results, and few have examined effects on colony development. Here we report the results of a comprehensive five-year study of the effects of the neonicotinoid imidacloprid on honey bee colonies. Colonies fed 5 ng/g (ppb) imidacloprid in sugar syrup showed increased brood production, lower temperature variability, higher CO 2 production and had more foragers compared to control colonies fed unadulterated syrup, but treatment did not affect adult bee numbers or average hive temperatures, and did not increase food stores, daily food acquisition or colony survivorship. These results suggest that imidacloprid contamination increased colony metabolism without improving colony productivity, and helps explain why some studies have reported no, or even positive, effects of neonicotinoids. Effect sizes were generally small but that could be attributed at least in part to variability in uncontrolled factors such as weather. We provide an explanation for the diverse effects of pesticide contamination on honey bees, and an improved understanding of how colonies are impacted., (Published by Elsevier Inc.)

Central place foraging pollinators tend to develop multi-destination routes (traplines) to exploit patchily distributed plant resources. While the formation of traplines by individual pollinators has been studied in detail, how populations of foragers use resources in a common area is an open question, difficult to address experimentally. We explored conditions for the emergence of resource partitioning among traplining bees using agent-based models built from experimental data of bumblebees foraging on artificial flowers. In the models, bees learn to develop routes as a consequence of feedback loops that change their probabilities of moving between flowers. While a positive reinforcement of movements leading to rewarding flowers is sufficient for the emergence of resource partitioning when flowers are evenly distributed, the addition of a negative reinforcement of movements leading to unrewarding flowers is necessary when flowers are patchily distributed. In environments with more complex spatial structures, the negative experiences of individual bees on flowers favour spatial segregation and efficient collective foraging. Our study fills a major gap in modelling pollinator behaviour and constitutes a unique tool to guide future experimental programs., Competing Interests: The authors have declared that no competing interests exist.

The current decline of invertebrates worldwide is alarming. Several potential causes have been proposed but metal pollutants, while being widespread in the air, soils and water, have so far been largely overlooked. Here, we reviewed the results of 527 observations of the effects of arsenic, cadmium, lead and mercury on terrestrial invertebrates. These four well-studied metals are considered as priorities for public health and for which international regulatory guidelines exist. We found that they all significantly impact the physiology and behavior of invertebrates, even at levels below those recommended as 'safe' for humans. Our results call for a revision of the regulatory thresholds to better protect terrestrial invertebrates, which appear to be more sensitive to metal pollution than vertebrates. More fundamental research on a broader range of compounds and species is needed to improve international guidelines for metal pollutants, and to develop conservation plans to protect invertebrates and ecosystem services., Competing Interests: Declaration of competing interest The authors declare no conflict of interest., (Copyright © 2021 The Authors. Published by Elsevier B.V. All rights reserved.)

Environmental pollutants can exert sublethal deleterious effects on animals. These include disruption of cognitive functions underlying crucial behaviours. While agrochemicals have been identified as a major threat to pollinators, metal pollutants, which are often found in complex mixtures, have so far been overlooked. Here, we assessed the impact of acute exposure to field-realistic concentrations of three common metal pollutants, lead, copper and arsenic, and their combinations, on honey bee appetitive learning and memory. All treatments involving single metals slowed down learning and disrupted memory retrieval at 24 h. Combinations of these metals had additive negative effects on both processes, suggesting common pathways of toxicity. Our results highlight the need to further assess the risks of metal pollution on invertebrates., Competing Interests: Competing interests The authors declare no competing or financial interests., (© 2021. Published by The Company of Biologists Ltd.)

Pollutants can have severe detrimental effects on insects, even at sublethal doses, damaging developmental and cognitive processes involved in crucial behaviours. Agrochemicals have been identified as important causes of pollinator declines, but the impacts of other anthropogenic compounds, such as metallic trace elements in soils and waters, have received considerably less attention. Here, we exposed colonies of the European honey bee Apis mellifera to chronic field-realistic concentrations of lead in food and demonstrated that consumption of this trace element impaired bee cognition and morphological development. Honey bees exposed to the highest of these low concentrations had reduced olfactory learning performances. These honey bees also developed smaller heads, which may have constrained their cognitive functions as we show a general relationship between head size and learning performance. Our results demonstrate that lead pollutants, even at trace levels, can have dramatic effects on honey bee cognitive abilities, potentially altering key colony functions and the pollination service., (Copyright © 2021 The Authors. Published by Elsevier Inc. All rights reserved.)

We examined how bees solve a visual discrimination task with stimuli commonly used in numerical cognition studies. Bees performed well on the task, but additional tests showed that they had learned continuous (non-numerical) cues. A network model using biologically plausible visual feature filtering and a simple associative rule was capable of learning the task using only continuous cues inherent in the training stimuli, with no numerical processing. This model was also able to reproduce behaviours that have been considered in other studies indicative of numerical cognition. Our results support the idea that a sense of magnitude may be more primitive and basic than a sense of number. Our findings highlight how problematic inadvertent continuous cues can be for studies of numerical cognition. This remains a deep issue within the field that requires increased vigilance and cleverness from the experimenter. We suggest ways of better assessing numerical cognition in non-speaking animals, including assessing the use of all alternative cues in one test, using cross-modal cues, analysing behavioural responses to detect underlying strategies, and finding the neural substrate.

Queens and workers are very distinct phenotypes that develop from the same genome. Larvae from worker cells up to 3.5 d old can be transferred to larger queen cells and will subsequently be reared as queens and develop into functional queens. This has become a very popular queen rearing practice in contemporary apiculture. Here we used RNA-Seq to study the consequences of rearing queens from transplanted worker larvae on the transcriptome of the adult queens. We found that queens reared from transferred older larvae developed slower, weighted less, and had fewer ovarioles than queens reared from transferred eggs, indicating queens were cryptically intercaste. RNA-Seq analysis revealed differentially expressed genes between queens reared from transferred larvae compared with queens reared from transferred eggs: the older the larvae transferred, the greater the number of differentially expressed genes. Many of the differentially expressed genes had functions related to reproduction, longevity, immunity, or metabolism, suggesting that the health and long-term viability of queens was compromised. Our finds verify the previous studies that adult queens reared from older transferred larvae were of lower quality than queens reared from transferred eggs or younger larvae., (© The Author(s) 2020. Published by Oxford University Press on behalf of Entomological Society of America. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Whether a female honey bee (Apis mellifera) develops into a worker or a queen depends on her nutrition during development, which changes the epigenome to alter the developmental trajectory. Beekeepers typically exploit this developmental plasticity to produce queen bee by transplanting worker larvae into queen cells to be reared as queens, thus redirecting a worker developmental pathway to a queen developmental pathway. We studied the consequences of this manipulation for the queen phenotype and methylome over four generations. Queens reared from worker larvae consistently had fewer ovarioles than queens reared from eggs. Over four generations the methylomes of lines of queens reared from eggs and worker larvae diverged, accumulating increasing differences in exons of genes related to caste differentiation, growth and immunity. We discuss the consequences of these cryptic changes to the honey bee epigenome for the health and viability of honey bee stocks., (Copyright © 2020. Published by Elsevier Ltd.)

Honeybees forage on diverse flowers which vary in the amount and type of rewards they offer, and bees are challenged with maximizing the resources they gather for their colony. That bees are effective foragers is clear, but how bees solve this type of complex multi-choice task is unknown. Here, we set bees a five-comparison choice task in which five colours differed in their probability of offering reward and punishment. The colours were ranked such that high ranked colours were more likely to offer reward, and the ranking was unambiguous. Bees' choices in unrewarded tests matched their individual experiences of reward and punishment of each colour, indicating bees solved this test not by comparing or ranking colours but by basing their colour choices on their history of reinforcement for each colour. Computational modelling suggests a structure like the honeybee mushroom body with reinforcement-related plasticity at both input and output can be sufficient for this cognitive strategy. We discuss how probability matching enables effective choices to be made without a need to compare any stimuli directly, and the use and limitations of this simple cognitive strategy for foraging animals.

Prior to leaving home, insects acquire visual landmark information through a series of well-choreographed walks or flights of learning [1-4]. This information allows them to pinpoint goals both when in their vicinity [5-7] and from locations they have not previously visited [8-10]. It is presumed that animals returning home match memorized views to their current view for successful view-based navigation [11]. While view-based navigation strategies have been incorporated into several navigation models [8, 12, 13], we still know little about how this behavior is performed by the insect brain. Mushroom bodies are essential for visual learning and memory [14-16], and therefore we investigated their role in view-based navigation in a visually oriented ant, Myrmecia midas. We injected the local anesthetic procaine [15, 17, 18] into the mushroom body vertical lobes (VLs) to selectively inhibit neural activity in this region. We compared the behavior of VL-procaine-treated ants with three groups: untreated control, VL-saline, and off-target (antennal lobe) procaine. Experienced foragers were collected, treated, and released in their familiar environment where we documented their behavior. Animals with procaine-inactivated VLs had tortuous paths and were unable to find their nest, whereas ants from the untreated and off-target procaine groups were well directed and were the most successful at returning home. Untreated animals walked faster when their gaze was directed toward home, and this behavior was eliminated by anesthetizing the VL region. Our data provide neurobiological evidence that the mushroom body vertical lobes are necessary for retrieving visual memories for successful view-based navigation., Competing Interests: Declaration of Interests The authors have no conflicts of interest to declare., (Copyright © 2020 Elsevier Inc. All rights reserved.)

Background: There is a dearth of literature describing factors supporting a successful transition from a career-development fellowship to resumption of the full complement of faculty roles., Purpose: Because little is known about the transition back to the full faculty role, a subset of Robert Wood Johnson Foundation (RWJF) Nurse Faculty Scholars (NFS) was interviewed to evaluate the self-identified challenges and opportunities that the scholars faced and factors contributing to their success when they reassumed the full faculty role., Methods: A subset of scholars from cohorts beginning the RWJF NFS program between 2008 and 2012 (n = 10) was interviewed by members of the NFS National Advisory Committee. NFSs identified challenges and opportunities faced as they transitioned to their faculty roles following completion of the career development fellowship as well as the character of support received from individuals in their organizational influencing the experience of the faculty transition. Evaluation outcomes include recommendations for transition planning for home institutions, colleagues and fellows., Results: NFSs identified transition challenges including managing multiple responsibilities and increased teaching demands coupled with loss of protected time and funding for scholarly work. Opportunities for career advancement were influenced by effective mentorship, institutional supports including advocacy and allocation of time and responsibilities consistent with continued research productivity. Issues contributing to a more difficult transition included non-supportive relationships among administrators and colleagues and newly assigned responsibilities that detracted from success in meeting expectations for tenure and promotion. Effective transition from fellow to faculty included plans for continued mentorship and stakeholder engagement of administration, mentors and faculty colleagues., Conclusions: Effective transition from fellow to the full complement of the faculty role benefits both the home institution and scholar. Positive outcomes may be contingent on scholar support and organizational investment during the transition period., (Copyright © 2020 Elsevier Inc. All rights reserved.)

For the materialist, the hard problem is fundamentally an explanatory problem. Solving it requires explaining why the relationship between brain and experience is the way it is and not some other way. We use the tools of the interventionist theory of explanation to show how a systematic experimental project could help move beyond the hard problem. Key to this project is the development of second-order interventions and invariant generalizations. Such interventions played a crucial scientific role in untangling other scientific mysteries, and we suggest that the same will be true of consciousness. We further suggest that the capacity for safe and reliable self -intervention will play a key role in overcoming both the hard and meta-problems of consciousness. Finally, we evaluate current strategies for intervention, with an eye to how they might be improved., (© The Author(s) 2020. Published by Oxford University Press.)

Human same-sex sexual attraction (SSSA) has long been considered to be an evolutionary puzzle. The trait is clearly biological: it is widespread and has a strong additive genetic basis, but how SSSA has evolved remains a subject of debate. Of itself, homosexual sexual behavior will not yield offspring, and consequently individuals expressing strong SSSA that are mostly or exclusively homosexual are presumed to have lower fitness and reproductive success. How then did the trait evolve, and how is it maintained in populations? Here we develop a novel argument for the evolution of SSSA that focuses on the likely adaptive social consequences of SSSA. We argue that same sex sexual attraction evolved as just one of a suite of traits responding to strong selection for ease of social integration or prosocial behavior. A strong driver of recent human behavioral evolution has been selection for reduced reactive aggression, increased social affiliation, social communication, and ease of social integration. In many prosocial mammals sex has adopted new social functions in contexts of social bonding, social reinforcement, appeasement, and play. We argue that for humans the social functions and benefits of sex apply to same-sex sexual behavior as well as heterosexual behavior. As a consequence we propose a degree of SSSA, was selected for in recent human evolution for its non-conceptive social benefits. We discuss how this hypothesis provides a better explanation for human sexual attractions and behavior than theories that invoke sexual inversion or single-locus genetic models., (Copyright © 2020 Barron and Hare.)

Pesticide residues have been linked to reduced bee health and increased honey bee colony failure. Most research to date has investigated the role of pesticides on individual honey bees, and it is still unclear how trace levels of pesticides change colony viability and productivity over seasonal time scales. To address this question we exposed standard bee colonies to chemical stressors known to have negative effects on individual bees, and measured the productivity of bee colonies across a whole year in two environments: near Tucson Arizona and Sydney Australia. We exposed hives to a trace amount of the neonicotinoid imidacloprid and to the acaricide thymol, and measured capped brood, bee and honey production, as well as the temperature and foraging force of the colonies. The effect of imidacloprid on colony dynamics differed between the two environments. In Tucson we recorded a positive effect of imidacloprid treatment on bee and brood numbers. Thymol was associated with short-term negative effects on bee numbers at both locations, and may have affected colony survival at one location. The overall benefits of thymol for the colonies were unclear. We conclude that long-term and colony-level measures of the effects of agrochemicals are needed to properly understand risks to bees., (Copyright © 2019. Published by Elsevier B.V.)

There is increasing worldwide concern about the impacts of pesticide residues on honey bees and bee colony survival, but how sublethal effects of pesticides on bees might cause colony failure remains highly controversial, with field data giving very mixed results. To explore how trace levels of the neonicotinoid pesticide imidacloprid impacted colony foraging performance, we equipped bees with RFID tags that allowed us to track their lifetime flight behavior. One group of bees was exposed to a trace concentration (5 μg/kg, ppb) of imidacloprid in sugar syrup while in the larval stage. The imidacloprid residues caused bees to start foraging when younger as adults and perform fewer orientation flights, and reduced their lifetime foraging flights by 28%. The magnitude of the effects of a trace imidacloprid concentration delivered only during larval stage highlights the severity of pesticide residues for bee foraging performance. Our data suggest that neonicotinoids could impact colony function by imbalancing the normal age based division of labor in a colony and reducing foraging efficiency. Understanding this mechanism will help the development of interventions to safeguard bee colony health.

Influences from the mother on offspring phenotype, known as maternal effects, are an important cause of adaptive phenotypic plasticity [1, 2]. Eusocial insects show dramatic phenotypic plasticity with morphologically distinct reproductive (queen) and worker castes [3, 4]. The dominant paradigm for honeybees (Apis mellifera) is that castes are environmentally rather than genetically determined, with the environment and diet of young larvae causing caste differentiation [5-9]. A role for maternal effects has not been considered, but here we show that egg size also influences queen development. Queens laid significantly bigger eggs in the larger queen cells than in the worker cells. Eggs laid in queen cells (QE), laid in worker cells (WE), and 2-day old larvae from worker cells (2L) were transferred to artificial queen cells to be reared as queens in a standardized environment. Newly emerged adult queens from QE were heavier than those from the other two groups and had more ovarioles, indicating a consequence of egg size for adult queen morphology. Gene expression analyses identified several significantly differentially expressed genes between newly emerged queens from QE and those from the other groups. These included a disproportionate number of genes involved in hormonal signaling, body development, and immune pathways, which are key traits differing between queens and workers. That egg size influences emerging queen morphology and physiology and that queens lay larger eggs in queen cells demonstrate both a maternal effect on the expression of the queen phenotype and a more active role for the queen in gyne production than has been realized previously., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

Honeybees (Apis mellifera) have haplodiploid sex determination: males develop from unfertilized eggs and females develop from fertilized ones. The differences in larval food also determine the development of females. Here we compared the total somatic gene expression profiles of 2-day and 4-day-old drone, queen and worker larvae by RNA-Seq. The results from a co-expression network analysis on all expressed genes showed that 2-day-old drone and worker larvae were closer in gene expression profiles than 2-day-old queen larvae. This indicated that for young larvae (2-day-old) environmental factors such as larval diet have a greater effect on gene expression profiles than ploidy or sex determination. Drones had the most distinct gene expression profiles at the 4-day larval stage, suggesting that haploidy, or sex dramatically affects the gene expression of honeybee larvae. Drone larvae showed fewer differences in gene expression profiles at the 2-day and 4-day time points than the worker and queen larval comparisons (598 against 1190 and 1181), suggesting a different pattern of gene expression regulation during the larval development of haploid males compared to diploid females. This study indicates that early in development the queen caste has the most distinct gene expression profile, perhaps reflecting the very rapid growth and morphological specialization of this caste compared to workers and drones. Later in development the haploid male drones have the most distinct gene expression profile, perhaps reflecting the influence of ploidy or sex determination on gene expression., (© 2017 Institute of Zoology, Chinese Academy of Sciences.)

Background: How to be an effective mentor is typically not taught formally because good mentoring is thought to beget good mentoring, but there is little concrete data to support that connection., Purpose: Scholars in the Robert Wood Johnson Foundation's Nurse Faculty Scholars (NFS) program were surveyed to find out if the mentoring they received influenced their subsequent mentoring., Method: The qualities that form the Mentorship Effectiveness Scale were used to investigate if the experience changed scholars' views of mentoring; open-ended questions provided an opportunity for scholars to describe additional insights., Results: Thirty-nine out of 93 scholars (42%) replied. Scholars were influenced by the mentoring they received: they were now more inclined to give guidance on professional issues, provide constructive critiques, suggest resources, acknowledge mentees' contributions, and challenge mentees to extend their abilities. Scholars' experiences made them more aware of the power of mentoring, provided them with insights into how they might do the same, convinced them that mentoring shouldn't be confined to one advisor, made them realize that one style of mentoring doesn't work for everyone, and improved their likelihood of using an individual development plan when they mentored., Conclusions: Effective mentoring does seem to beget effective mentoring because the interaction with mentors seems to leave a lasting impression and the perceived obligation to "pay it forward.", (Copyright © 2018 Elsevier Inc. All rights reserved.)

Honey bee foragers must supply their colony with a balance of pollen and nectar to sustain optimal colony development. Inter-individual behavioural variability among foragers is observed in terms of activity levels and nectar vs. pollen collection, however the causes of such variation are still open questions. Here we explored the relationship between foraging activity and foraging performance in honey bees (Apis mellifera) by using an automated behaviour monitoring system to record mass on departing the hive, trip duration, presence of pollen on the hind legs and mass upon return to the hive, during the lifelong foraging career of individual bees. In our colonies, only a subset of foragers collected pollen, and no bee exclusively foraged for pollen. A minority of very active bees (19% of the foragers) performed 50% of the colony's total foraging trips, contributing to both pollen and nectar collection. Foraging performance (amount and rate of food collection) depended on bees' individual experience (amount of foraging trips completed). We argue that this reveals an important vulnerability for these social bees since environmental stressors that alter the activity and reduce the lifespan of foragers may prevent bees ever achieving maximal performance, thereby seriously compromising the effectiveness of the colony foraging force.

Biogenic amines modulate a range of social behaviours, including sociability and mechanisms of group cohesion, in both vertebrates and invertebrates. Here, we tested if the biogenic amines modulate honey bee (Apis mellifera) sociability and nestmate affiliation. We examined the consequences of treatments with biogenic amines, agonists and antagonists on a bee's approach to, and subsequent social interactions with, conspecifics in both naturally hive-reared bees and isolated bees. We used two different treatment methods. Bees were first treated topically with compounds dissolved in the solvent dimethylformamide (dMF) applied to the dorsal thorax, but dMF had a significant effect on the locomotion and behaviour of the bees during the behavioural test that interfered with their social responses. Our second method used microinjection to deliver biogenic amines to the head capsule via the ocellar tract. Microinjection of dopamine and a dopamine antagonist had strong effects on bee sociability, likelihood of interaction with bees, and nestmate affiliation. Octopamine treatment reduced social interaction with other bees, and serotonin increased the likelihood of social interactions. HPLC measurements showed that isolation reduced brain levels of biogenic amines compared to hive-reared bees. Our findings suggest that dopamine is an important neurochemical component of social motivation in bees. This finding advances a comparative understanding of the processes of social evolution., Competing Interests: The authors have declared that no competing interests exist.

Precise, objective data on brood and honey levels in honey bee colonies can be obtained through the analysis of hive frame photographs. However, accurate analysis of all the frame photographs from medium- to large-scale experiments is time-consuming. This limits the number of hives than can be practically included in honeybee studies. Faster estimation methods exist but they significantly decrease precision and their use requires a larger sample size to maintain statistical power. To resolve this issue, we created 'CombCount' a python program that automatically detects uncapped cells to speed up measurements of capped brood and capped honey on photos of frames. CombCount does not require programming skills, it was designed to facilitate colony-level research in honeybees and to provide a fast, free, and accurate alternative to older methods based on visual estimations. Six observers measured the same photos of thirty different frames both with CombCount and by manually outlining the entire capped areas with ImageJ. The results obtained were highly similar between both the observers and the two methods, but measurements with CombCount were 3.2 times faster than with ImageJ (4 and 13 min per side of the frame, respectively) and all observers were faster when using CombCount rather than ImageJ. CombCount was used to measure the proportions of capped brood and capped honey on each frame of 16 hives over a year as they developed from packages to full-size colonies over about 60 days. Our data describe the formation of brood and honey stores during the establishment of a new colony., Competing Interests: The authors have declared that no competing interests exist.

The capacity to learn abstract concepts such as 'sameness' and 'difference' is considered a higher-order cognitive function, typically thought to be dependent on top-down neocortical processing. It is therefore surprising that honey bees apparantly have this capacity. Here we report a model of the structures of the honey bee brain that can learn sameness and difference, as well as a range of complex and simple associative learning tasks. Our model is constrained by the known connections and properties of the mushroom body, including the protocerebral tract, and provides a good fit to the learning rates and performances of real bees in all tasks, including learning sameness and difference. The model proposes a novel mechanism for learning the abstract concepts of 'sameness' and 'difference' that is compatible with the insect brain, and is not dependent on top-down or executive control processing., Competing Interests: The authors have declared that no competing interests exist.

Pesticides are considered one of the major contemporary stressors of honey bee health. In this study, the effects of short-term exposure to lambda-cyhalothrin on lifespan, learning, and memory-related characteristics of Apis mellifera were systematically examined. Short-term exposure to lambda-cyhalothrin in worker bees reduced lifespan, affected learning and memory performance, reduced the homing ability, and influenced the expression levels of two learning and memory-related genes of A. mellifera. This research identifies the nature of the sublethal effects of lambda-cyhalothrin on bees and the level of exposure that can be harmful to bee health. This new information will assist in establishing guidelines for the safe use of lambda-cyhalothrin in the field.

Patterns in within-day hive weight data from two independent datasets in Arizona and California were modeled using piecewise regression, and analyzed with respect to honey bee colony behavior and landscape effects. The regression analysis yielded information on the start and finish of a colony's daily activity cycle, hive weight change at night, hive weight loss due to departing foragers and weight gain due to returning foragers. Assumptions about the meaning of the timing and size of the morning weight changes were tested in a third study by delaying the forager departure times from one to three hours using screen entrance gates. A regression of planned vs. observed departure delays showed that the initial hive weight loss around dawn was largely due to foragers. In a similar experiment in Australia, hive weight loss due to departing foragers in the morning was correlated with net bee traffic (difference between the number of departing bees and the number of arriving bees) and from those data the payload of the arriving bees was estimated to be 0.02 g. The piecewise regression approach was then used to analyze a fifth study involving hives with and without access to natural forage. The analysis showed that, during a commercial pollination event, hives with previous access to forage had a significantly higher rate of weight gain as the foragers returned in the afternoon, and, in the weeks after the pollination event, a significantly higher rate of weight loss in the morning, as foragers departed. This combination of continuous weight data and piecewise regression proved effective in detecting treatment differences in foraging activity that other methods failed to detect., Competing Interests: The authors have declared that no competing interests exist.

Brain structure and learning capacities both vary with experience, but the mechanistic link between them is unclear. Here, we investigated whether experience-dependent variability in learning performance can be explained by neuroplasticity in foraging honey bees. The mushroom bodies (MBs) are a brain center necessary for ambiguous olfactory learning tasks such as reversal learning. Using radio frequency identification technology, we assessed the effects of natural variation in foraging activity, and the age when first foraging, on both performance in reversal learning and on synaptic connectivity in the MBs. We found that reversal learning performance improved at foraging onset and could decline with greater foraging experience. If bees started foraging before the normal age, as a result of a stress applied to the colony, the decline in learning performance with foraging experience was more severe. Analyses of brain structure in the same bees showed that the total number of synaptic boutons at the MB input decreased when bees started foraging, and then increased with greater foraging intensity. At foraging onset MB structure is therefore optimized for bees to update learned information, but optimization of MB connectivity deteriorates with foraging effort. In a computational model of the MBs sparser coding of information at the MB input improved reversal learning performance. We propose, therefore, a plausible mechanistic relationship between experience, neuroplasticity, and cognitive performance in a natural and ecological context.

Foraging in honeybees is energetically demanding. Here, we examined whether stressors, which generally increase metabolic demands, can impair foraging performance. A controlled non-pathogenic stressor (immune challenge) resulted in a change in the foraging preferences of bees. It reduced pollen foraging and increased the duration of trips in pollen foragers. Stress also reduced the amount of octopamine in the brain of pollen foragers (a biogenic amine involved in the regulation of foraging and flight behaviour in insects). According to the literature, flight metabolic rate is higher during pollen foraging than during nectar foraging, and nectar gives a higher energetic return relative to the foraging effort when compared with pollen. We thus propose that stress might be particularly detrimental to the performance of pollen foragers, and stressed bees prefer the energy-rich resource of nectar. In conclusion, stress, even at low levels, could have consequences for bee foraging behaviour and thereby the nutritional balance of the colony., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

Underpinning the formation of a social group is the motivation of individuals to aggregate and interact with conspecifics, termed sociability. Here, we developed an assay, inspired by vertebrate approaches to evaluate social behaviours, to simultaneously examine the development of honey bee ( Apis mellifera ) sociability and nestmate affiliation. Focal bees were placed in a testing chamber which was separated from groups of nestmates and conspecific non-nestmates by single-layer mesh screens. Assessing how much time bees spent contacting the two mesh screens allowed us to quantify simultaneously how much bees sought proximity and interaction with other bees and their preference for nestmates over non-nestmates. Both sociability and nestmate affiliation could be detected soon after emergence as an adult. Isolation early in adult life impaired honey bee sociability but there was no evidence for a critical period for the development of the trait, as isolated bees exposed to their hive for 24 h when as old as 6 days still recovered high levels of sociability. Our data show that, even for advanced social insects, sociability is a developmental phenomenon and experience dependent., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2018. Published by The Company of Biologists Ltd.)

Drug addiction is a chronic relapsing behavioral disorder. The high relapse rate has often been attributed to the perseverance of drug-associated memories due to high incentive salience of stimuli learnt under the influence of drugs. Drug addiction has also been interpreted as a memory disorder since drug associated memories are unusually enduring and some drugs, such as cocaine, interfere with neuroepigenetic machinery known to be involved in memory processing. Here we used the honey bee (an established invertebrate model for epigenomics and behavioral studies) to examine whether or not cocaine affects memory processing independently of its effect on incentive salience. Using the proboscis extension reflex training paradigm we found that cocaine strongly impairs consolidation of extinction memory. Based on correlation between the observed effect of cocaine on learning and expression of epigenetic processes, we propose that cocaine interferes with memory processing independently of incentive salience by directly altering DNA methylation dynamics. Our findings emphasize the impact of cocaine on memory systems, with relevance for understanding how cocaine can have such an enduring impact on behavior.

The defence of a society often requires that some specialized members coordinate to repel a threat at personal risk. This is especially true for honey bee guards, which defend the hive and may sacrifice their lives upon stinging. Central to this cooperative defensive response is the sting alarm pheromone, which has isoamyl acetate (IAA) as its main component. Although this defensive behaviour has been well described, the neural mechanisms triggered by IAA to coordinate stinging have long remained unknown. Here we show that IAA upregulates brain levels of serotonin and dopamine, thereby increasing the likelihood of an individual bee to attack and sting. Pharmacological enhancement of the levels of both amines induces higher defensive responsiveness, while decreasing them via antagonists decreases stinging. Our results thus uncover the neural mechanism by which an alarm pheromone recruits individuals to attack and repel a threat, and suggest that the alarm pheromone of honey bees acts on their response threshold rather than as a direct trigger., (© 2018 The Author(s).)

Honey bee dance has been intensively studied as a communication system, and yet we still know very little about the neurobiological mechanisms supporting how dances are produced and interpreted. Here, we discuss how new information on the functions of the central complex (CX) of the insect brain might shed some light on possible neural mechanisms of dance behaviour. We summarise the features of dance communication across the species of the genus Apis We then propose that neural mechanisms of orientation and spatial processing found to be supported by the CX may function in dance communication also, and that this mechanistic link could explain some specific features of the dance form. This is purely a hypothesis, but in proposing this hypothesis, and how it might be investigated, we hope to stimulate new mechanistic analyses of dance communication., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2017. Published by The Company of Biologists Ltd.)