Does The Honeybee Change The Concentration Of Nectar While En Route To The Hive

Abstract

Honey bee foragers must supply their colony with a balance of pollen and nectar to sustain optimal colony evolution. Inter-individual behavioural variability among foragers is observed in terms of activity levels and nectar vs. pollen drove, however the causes of such variation are still open up questions. Here we explored the relationship between foraging activity and foraging performance in love bees (Apis mellifera) past using an automated behaviour monitoring system to record mass on parting the hive, trip duration, presence of pollen on the hind legs and mass upon render to the hive, during the lifelong foraging career of private bees. In our colonies, just a subset of foragers collected pollen, and no bee exclusively foraged for pollen. A minority of very agile bees (19% of the foragers) performed fifty% of the colony's total foraging trips, contributing to both pollen and nectar drove. Foraging operation (amount and rate of food drove) depended on bees' private feel (amount of foraging trips completed). We argue that this reveals an of import vulnerability for these social bees since environmental stressors that modify the action and reduce the lifespan of foragers may prevent bees ever achieving maximal functioning, thereby seriously compromising the effectiveness of the colony foraging forcefulness.

Introduction

Social insects are reliant on the forager caste to supply resources for the whole colony^ane. However, not all foragers contribute equally to the collective try of colony provisioning^ii,3,4, which raises the questions of how and why such inter-private variability is observed.

In social bees, adequate colony diet requires a supply of both pollen (rich in proteins and fat) and nectar (elementary carbohydrates)⁵. It has been argued that having different individuals specialised for nectar or pollen collection is the most efficient strategy at the colony level⁶, due to the unlike spatio-temporal distributions of these major nutritional resource in the field and to the need for specific behavioural skills to collect each of them. This is generally causeless to be true for dear bees where pollen and nectar foragers are considered as different behavioural castes^{7,8,9,ten,eleven} that differ in their brain neuropeptide profiles¹², sucrose response threshold¹³, ovary size¹⁴, levels of vitellogenin (a yolk precursor protein)¹⁵ and responses to social stimuli (pollen foragers rely more on dance communication)¹⁶. Together with evidence for genetic variation^eight,17,18, these observations advise that pollen and nectar collection are evolved specialisations inside the foraging forcefulness of the colony^17,19,twenty. Yet, recent behavioural studies suggest that the distinction between pollen and nectar foraging may not be accented. A proportion of the foragers seem to collect both resources, or may change specialisation as they age^21,22. These studies, either conducted on a very express number of individuals (less than thirty bees)²²,or in spatially restricted flight cages with a few abundant artificial pollen or nectar sources²¹, telephone call for further investigations. And so far, no report has analysed the long-term foraging preferences of a large cohort of honey bees in the natural environment, and thus, data on how foragers partition their effort between pollen and nectar drove across their lifetime are express.

In improver to variation in the blazon of resource collected by foragers^{7,8,9,10,11,12,13}, some individuals provender more actively than others^21,23. Tenczar et al.²¹ reported that the bulk of foraging trips were performed by a minority of honey bee foragers, which they termed "elite bees". In this study, however, the authors were not able to measure out bees' foraging performance in terms of amounts or rates of nectar or pollen collection, but but on the number of trips completed. Without this data, it is not possible to know if the most agile foragers are also the most successful. One potential cause of inter-individual variation in foraging performance is the amount of experience gained by foragers over previous trips. Dukas²² and Schippers et al.²⁴ both showed that individual honey bees improve their foraging performance with experience. While both studies recorded individual behaviour in detail across the whole foraging career of individuals bees, they did then on a rather limited sample size and focused on nectar resources only.

Hither, we explored the nature and possible causes of variation in foraging activity and performance between honey bee (Apis mellifera) foragers using an automated behavioural tracking system at the hive entrance to monitor the foraging behaviour of a big number of bees (>260) from two colonies in the natural surround (Fig. 1). The hives were located in a room and connected to the outside surroundings via a specially designed archway containing baffles that forced bees to go out the hive forth i path and to enter using a dissimilar path. Our system employed radio frequency identification (RFID) engineering for the detection of arrival and departure times of private bees at the colony entrance^{25,26,27,28,29,30,31} and interpolation of individuals' foraging trips²⁵. We also used a digital camera to photograph returning foragers and digital balances to tape their mass. From these data, nosotros analysed how individual bees differed in foraging performance throughout their entire foraging career.

Results

Bees varied in their tendency to collect pollen and non-pollen resources

Nosotros analysed the foraging activeness of 564 bees for which we had information near the type of resource nerveless for at to the lowest degree one trip (Table 1). These bees performed an average of 19 foraging trips in their lifetime (mean ± SE, colony i: 17 ± 1 trips, Due north = 295; colony 2: 21 ± 1 trips, N = 269) and their foraging span was less than a calendar week (colony 1: 4.20 ± 0.18 days, N = 295; colony ii: 4.85 ± 0.xviii days, N = 269).

Table 1 Number of foragers and foraging trips recorded per colony.

Full size tabular array

On boilerplate 27% of bees foraged at to the lowest degree in one case for pollen (29% for colony 1, and 25% for colony 2, Table 1). None of these bees foraged exclusively on pollen. All of them performed a combination of pollen trips (cases when bees returned with pollen load) and not-pollen trips (cases when bees returned with no pollen load). We considered them as "mixed foragers". Non-pollen trips could be unsuccessful trips, or trips for nectar or water. Incidences of pollen drove were distributed throughout the experiment and foragers varied in the number of trips equally well every bit in the percent of pollen trips they performed on different days (Fig. 2A).

For non-pollen trips the difference in mass on departure and render to the colony was either positive of effectually nil, and thus provides information near the amount of resource collected (mean ± SE, =−0.56 ± 0.24 mg, Northward = 925). For pollen trips the hateful mass difference was negative (mean ± SE, −1.54 ± 12.5 mg, N = 136) despite bees returning with big and obvious pollen loads. From this, we infer that foragers returning with pollen must have expended mass (which we presume to be crop contents) during their pollen collection trip. Consequently, we could not employ mass to measure the corporeality of pollen collected. Mixed foragers were heavier when leaving the hive for a pollen collecting trip than when leaving the hive for a non-pollen trip (Tabular array 2b).

Table 2 Foraging strategies of mixed foragers.

Total size table

Bees improved their foraging operation and likelihood of pollen collection, with feel

For each bee, we found a quadratic relationship betwixt the probability of collecting pollen on a foraging trip and the number of foraging trips already performed (LRTest: Chisq = 56.324, df = 1, P < 0.001). This indicates that bees tended to increase the collection of pollen during their first flights up to a maximum where the probability of pollen collection thereafter decreased with boosted foraging experience (Fig. 2A, Wald ChiSq Test for Binomial GLMM; linear predictor Chisq_i,2.215 = 97.07, P < 0.001; quadratic predictor: Chisq_1,2.215 = 34.72, P < 0.001). The probability of collecting pollen was similar betwixt the two colonies (Chisq_ane,2.215 = 3.03, P = 0.082). However, we found a stronger quadratic decline in the proportion of pollen collected with cumulative trips in colony 1 (tested during Australian autumn) than in colony 2 (tested in Australian bound) (Chisq_i,2.215 = 11.84, P < 0.001), suggesting that environmental weather condition influenced pollen availability and collection.

Foragers performed more trips per solar day until about day 9 after starting foraging (GLMM Poisson, Chisq_1,2.402 = ix.32, P = 0.002). From this point, the number of trips performed each day per bee stabilised (GLMM Poisson, Chisq_1,23 = 0.49, P = 0.484). Piecewise regression using the estimated break point at mean solar day 9 produced a meliorate fit than a regression using the complete dataset (RSS_piecewise = 256661, RSS_{complete_dataset} = 280596, F = 60.99, P < 0.001, Fig. 2B)

Bees showed an increase in performance (weight deviation between weight on inflow and weight on deviation) for not-pollen trips with successive trip number (Fig. 2C, LMM, F_(1,925) = 6.45, P = 0.016). A similar relationship was observed for foraging efficiency (mass difference between deviation and arrival divided past the trip duration, mg/min; Table S2, Fig. S2).

Well-nigh foraging trips were performed by a minority of elite bees

We examined the proportion of foraging trips performed by each bee, relative to all the foraging trips of their colony. Visualisation of the inter-individual variation in foraging attempt seen in each colony using a Lorenz curve³² indicates that around nineteen% of the tagged foragers performed 50% of the total number of trips recorded (17.29% for colony 1, 20.45% for colony 2; Fig. iii). Following Tenczar²¹, we describe these very agile bees as elite bees. Nosotros then examined the skew of foraging activity between all foragers past computing a Gini index³³ with values comprised between 0 (if all individuals contributed equally to the common job) and 1 (if only i individual performed the whole foraging task). Nosotros obtained a Gini index of 0.49 for colony 1 and 0.46 for colony two, meaning that in both colonies, not all individuals contributed equally to the common foraging job.

Elite bees performed better than non-aristocracy bees

When comparing the operation of elite and not-elite bees we found a larger proportion (66%) of mixed foragers within the elite bees than inside the non-elite bees (17.five%) (Fig. S1A, colony 1: 72% of aristocracy bees are mixed foragers, vs. 20% of non-aristocracy bees: of mixed foragers: χ² = 55.68, df = i, P < 0.0001; colony ii: 60% of aristocracy bees are mixed foragers vs. 15% of non-elite bees: χ² = 46.99, df = i, P < 0.0001).

For non-pollen trips, elite bees had a greater gain in mass during their foraging trip than not-elite bees, indicating that elite bees returned more resources to the colony per trip (Fig. S1B). The differences in functioning of elite and non-aristocracy bees could be explained by the differences in experience between these two groups. Models considering individual experience on foraging operation showed a meaning outcome of experience, but no significant consequence of elite bees as a fixed factor (Table S2).

Discussion

Nosotros combined RFID tracking with video and mass recordings to explore the nature of variation in foraging functioning between individual honey bees, and the possible causes of that variation. We found that foragers differed greatly in their lifetime foraging behaviour. Foraging performance and likelihood of collecting pollen increased as bees gained experience.

Experienced foragers were more likely to collect pollen

In our study, scoring foragers that returned with pollen loads was unambiguous, but for bees that lacked pollen loads the nature of their trip was unclear. For bees that returned to the hive without pollen, we could not discriminate between an unsuccessful trip, a successful nectar collection or a successful h2o drove, and thus we classified trips as simply pollen or not-pollen. No bee collected pollen exclusively, which was concordant with other observations of behavioural introgression between pollen and nectar specialisations^{thirteen,21,22,34,35,36,37}. In our data, nonetheless, it was striking that pollen collection was performed by a minority of individuals, and commonly only individuals that had accumulated substantial foraging experience. If these foragers are lost to the colony (due to predation, adverse conditions or exposure to pesticides that disorient them³¹) in that location could be severe consequences for the pollen supply to the colony.

Nosotros notation however that our experimental design was constrained by the use of colonies at different times of the year. The pass up of pollen drove observed in experienced foragers of colony 1 (tested during Australian autumn) suggests that pollen availability was lower at this fourth dimension of the year. In addition, nosotros used pocket-size colonies with a minimum amount of breed because of the limitations imposed by the RFID sensors on forager traffic at the entrance. Time to come experiments on full-size commercial honey bee colonies at different periods of the year will be needed to assess the extent to which this pattern is dependent on flavour and population size.

Elite bees undertook the majority of the foraging trips

All foragers tended to increase their number of foraging trips per day as they gained experience, until they reached a plateau of activity towards the end of their foraging career. Withal, all foragers did non contribute equally to the total foraging activity of their colony (Fig. 3). Rather we noticed a strong skew with a minority of very active bees (hither and previously referred to as the aristocracy bees²¹) undertaking the majority of the colony's nectar and pollen foraging trips. In our report 19% of the foragers completed more than 50% of the full number of foraging trips in their colonies. In the same species, Tenczar et al.²¹ compared private activeness with total colony activity on a twenty-four hours-by-day basis because of a limit in the accuracy of their RFID sensor detection. When applying this particular analysis to our data (Fig. S3), we found 12% of the workers performed more than than l% of the colony's daily activeness, which is within the same range to the values reported by Tenczar et al. (effectually 20%)²¹.

Elite bees performed meliorate because they were more than experienced

Prior social insect studies observing like skews in foraging activity either failed to identify whatsoever detail link between performance and action^21,23, or simply reported a positive correlation between activity and efficiency^38,39. In our data, nosotros found a articulate relationship between individual performance and experience (Figs 2 and S2). The simplest caption for this is that through experience bees larn to improve their foraging performance, which is consistent with suggestions from earlier studies^{22,24,twoscore,41}. How this occurs is unknown, but cerebral studies have shown bees can learn to improve their foraging skills through trial-and-error⁴¹, their navigation between known food sources^{42,43,44,45,46,47}, their discrimination of flowers (based on colours, shapes, odours or textures⁴⁸) and their flower treatment^49,fifty. Whatever or all of these factors could explicate the improvements in natural foraging performance with experience nosotros observed in this study. Hereafter experiments should also examination whether environmental complexity (resources distribution, foraging distances, visual cues) affects the relationship between foraging experience and foraging performance, since individuals may learn more while making fewer trips in environments with complex structures.

Implications for understanding the impact of stress on a colony

We showed that the most active and experienced bees are also the nearly efficient and high performing foragers in a colony. Tenczar et al.²¹ argued that elite bees can be rapidly replaced by a 'reserve' of less active foragers, but these authors did not examine individual foraging performance. Hither we demonstrated that performance and efficiency is caused through experience. Therefore, while the activeness of an elite bee can be taken over quickly by other foragers, these new foragers many not be as efficient until they take gained feel. Further, whatsoever stressors that shorten foragers' lifespan, such equally pesticides, pollutants, unbalanced diets, pathogens and viruses^{31,51,52,53,54,55,56,57,58,59,60,61}, may preclude bees from accumulating enough experience to maximise their foraging contribution to the colony, ultimately leading to the recruitment of poorly efficient, precocious, foragers⁶². These interactions betwixt stress, foraging force age composition, individual forager experience and performance deserve further study as they may help united states understand the process of colony failure.

Cloth and Methods

Experimental hive

Honey bees (Apis mellifera) were obtained from the research apiary of Macquarie University (Sydney, NSW, Commonwealth of australia). The experimental hive was a iv-frame nucleus hive (wooden box of 56 × 23 × 28 cm containing 4 standard Langstroth frames), placed in a dark room at the constant temperature of 24 °C. The hive independent two frames of honey and pollen, ane frame of capped breed and i frame of polystyrene to fill the remaining space in the hive box.

The hive was connected to the outside environment via a especially designed archway containing baffles that forced bees to go out the hive along 1 path and to enter using a different path (Fig. ane). Each path was made of transparent plastic tubes (one cm diameter) that passed beyond an RFID antenna (Invengo, Guangzhou, China) and a microbalance pan (A&D Visitor Ltd, Japan). To prevent bees entering or exiting through the wrong tube, nosotros attached inwardly tapering plastic beard at the end of each tube. Bees landing at the archway were funneled toward the single entrance tube.

Sections of the entry and get out tubes ran across dynamic micro balances of one mg sensitivity). The balances captured the mass of bees as they entered and exited the hive. Sometimes several bees crossed the balance at the same time and thus, the mass did not reverberate the mass of just one individual. For this reason, we retained simply values between 60 and 150 mg, which we evaluated equally a conservative realistic individual mass range for honey bees. Test of the videos of the entrance tunnel indicated that these limits were realistic.

Along the archway path bees also passed beneath a webcam (Logitech), placed in a plastic box illuminated with white LED light, in order to video record the archway tube. A motion detection video recording software (Netcam Studio X, Moonware Studios) was used to capture video footages of returning bees, thus assuasive a visual cess of the presence or absence of pollen in the corbiculae on the bees' legs).

Automatic gates (micro-controlled servos connected to infrared emitter/receiver) regulated the traffic of bees inside each path. The gates were placed at the commencement of the archway and exit tubes. When a bee walked through the tubes and broke the beam of an infrared emitter/receiver, the continued gate would close behind the bee for 10 seconds. This fourth dimension was an estimation of the maximum fourth dimension needed for a bee to cross the RFID antenna and the balance. The infrared beams and gates were all continued and monitored using a single-board microcontroller system (Arduino, Adafruit Industries, United states).

Experimental bees

This arroyo has been used to study bee behaviour for more than a decade²⁵ to examine the affect of environmental stressors on bee foraging behaviour^{26,27,28,29,xxx,31}, or to reveal individual foraging strategies^21,63. RFID systems utilise tags, each with a unique digital ID that can be attached to bees. RFID tags were obtained from Invengo (Guangzhou, China). Each circular tag had a diameter of four mm and a mass of 1 mg and could be stock-still to the bees' dorsal thorax with gum (Loctite, Gel Super Glue). Each RFID tag had a unique 12-byte hexadecimal identifier that allowed the states to track individual bees as they were detected past each antenna on exiting and inbound the hive.

The hive was established with about 500 background (untagged) bees of mixed ages and a queen taken from a single colony. Newly emerged bees carrying individually programmed RFID tags were successively added to this hive. To source newly emerged bees nosotros collected brood frames from upward to eight dissimilar colonies over the course of the 2 experimental replicates to provide a diverse source of brood for the experiment. For each brood collection, frames were collected from two or three of the eight colonies. Brood frames were stored overnight in an incubator maintained at 37 °C. The next morning RFID tags were glued to the thorax of the newly emerged bees.

Tagged bees were added to the colony at different times. Nosotros divers a accomplice every bit a group of bees tagged and added to the colony over up to four successive days. Each cohort came from a combination of different hives in order to increase multifariousness. Over the five weeks of the experiment, the hive received around 4,400 bees in total distributed inside three cohorts. Approximately 500 tagged bees were added every day for the first four days of the experiment (commencement cohort). From day 21 of the experiment a further 1,800 tagged bees were added over four days (450 bees a day, second cohort). From solar day 35 of the experiment a farther 600 bees were added over four days (150 bees a twenty-four hour period, third cohort).

The experiment was conducted twice: colony one, from Apr-May 2015 (Australian autumn); colony ii, from November-Dec 2015 (Australian spring). In colony 1, the queen died after two weeks and was replaced with a queen mandibular pheromone substitute (BeeBoost, Hornsby beekeeping supply, Australia).

Bees foraged in the surrounding suburban Australian environment including several nature reserves and private gardens. Such an environment provided nectar and pollen menses during the two seasons of the experiment, with a predominance of flowering native trees and bushes such as different eucalyptus species.

Collating data on bee trips

RFID data for each trip included the RFID identification for the individual, the date and time it left the hive, and the date and time of its return to the hive, thus enabling united states of america to calculate the duration of trips. Trips that lasted less than 10 southward were considered as non-foraging trips⁶² and removed from the dataset. RFID readings were fourth dimension-matched with readings from the balances to capture the mass of bees on departure and return to the colony. Departure in both mass measures gave an indication of resources collection (positive values) or expenditure (negative values). Videos taken upwardly to 20 due south before an entry RFID detection were inspected to score whether the tagged returning bee carried pollen (P) on its legs (run into Supplementary Materials Video S1 in Dataset S2), no pollen (NP, come across Supplementary Materials Video S2 in Dataset S2) or could non exist reliably scored (NA) (i.east. when the bee legs were occluded on the video due to body bending or to the simultaneous presence of other bees in the entrance tube). The maximum time for a bee to travel from the webcam to the RFID antenna was visually assessed as 20 southward.

Data reliability

A total of 8,640 bees were tagged during the two replicates of the experiment (iv,390 for colony 1 and 4,250 for colony 2). From the initial dataset, we excluded bees that performed only one trip and bees that had only 'NA' as load type over all their trips. In the final dataset, three,432 bees (one,728 for colony 1 and ane,704 for colony 2) were retained. We speculate that the discrepancy between last bee counts and initial bees tagged is due to many bees losing their tags inside the hive, some of the tags being damaged during the tagging process and rendered unreadable, and some bees non returning to the hive during their first trip, for reasons of health or inability to wing due to poor positioning of the tag. For each bee, we excluded the first v trips, which are more probable orientation flights than foraging flights⁶². A summary of our trip dataset is given in Table 1. The loftier number of 'NA' trips was mainly due to photographic camera software issues, which was unfortunately virtually problematic for colony 2. The complete final information set up is provided in Dataset S1.

Information analyses

Information were analysed in R version three.2.3 (operating via Rstudio, version 1.0.136⁶⁴).

Differences between aristocracy and non-elite foragers, in age at first foraging trip and in average mass on departure, were tested with a Wilcoxon rank sum test. Differences in the proportions of mixed foragers in elite and non-elite bees were tested with a Chi-square test.

The correlation between the average mass difference for not-pollen trips and bee activeness was tested with a linear mixed model (LMM) using the function "lmer" in the package lme4 ⁶⁵ (p-values were extracted using the bundle lmerTest ⁶⁶).

The correlation betwixt the total mass differences for not-pollen trips and bee activity was tested using linear and quadratic mixed models, with "lmer". Nosotros verified the existence of a quadratic relationship past comparing the linear and the quadratic models using a Loglikelihood Ratio Examination (LRTest⁶⁷). If the p-value was less than 0.05, we kept the model with the lower log-likelihood value.

Variation in mass divergence for not-pollen trips according to the number of trips was tested using linear and logarithmic models with "lmer". Nosotros verified the better fitting of the logarithmic relationship using the LRTest.

Variation in the number of foraging trips per mean solar day according to the number of foraging days were tested using a GLMM 4 count data with Poisson distribution error using the function "glmer" in lme4. Afterwards visual inspection the trend between the number of foraging trips every bit a function of foraging days revealed an apparent stabilization of foraging trips effectually after 9 days of experience. To examination whether this stabilization was statistically significant, we fitted a piecewise mixed model splitting the dataset into two unlike regressions which break point was determined using the function "segmented" in the package segmented ⁶⁸. We compared the fits of the models (piecewise vs complete dataset GLMM) using the Balance Sum of Square (RSS) analysis. The lower the RSS the ameliorate the fit.

We estimated the probability of foraging for pollen according to experience (number of foraging trips) using a GLMM with Binomial distribution error, using the function "glmer". We verified the existence of a quadratic human relationship using the LRTest. Differences in mass at departure for non-pollen trips between not-pollen foragers (bees that did not perform any pollen trip) and mixed foragers (bees that nerveless pollen at to the lowest degree once) were analysed using a GLMM with Poisson distribution error.

For all models, colony identity was included equally a covariate and bee identity nested in cohort identity was included as a random factor. Day identity was also included every bit a random factor to control for ecology variation betwixt days. The colony of origin of the bees was found to have no event when included every bit a fixed factor in the models. Nosotros thus did non include colony of origin as a potential explanatory variable in our analyses.

All minimum adequate models were selected by comparing their Akaike Information Criterion (AIC) to zilch models⁶⁹ based on an analysis of variance (F test) in example of linear mixed model and analysis of deviance (Chisq test) in example of a generalized linear mixed model using respectively the function "anova" of the stats parcel and the role "Anova" of the automobile package (see details in Tabular array S2). All the analyses of variance produced with the parcel lmerTest were run using the default Satterthwaite caste of freedom approximation⁷⁰.

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Acknowledgements

We give thanks Andrew Allen for his advices on statistical analyses. S.K. was funded by a PhD fellowship from the French Ministry of Research and MQRS scholarship from Macquarie University. J.Thousand.D. was funded by the Agence Nationale de la Recherche (ANR-13-ADAP-0002). A.B.B. was funded by the Australian Inquiry Council (ARC Future Fellowship no. 140100452) and the United states of america Department of Agriculture ARS agreement no: 58-5342-3-004F. C.P. and Chiliad.L. were funded by the CNRS and a grant of the Agence Nationale de la Recherche to ML (ANR-xvi-CE02-0002-01).

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Author notes

Andrew B. Barron and Mathieu Lihoreau contributed as.

Affiliations

Research Centre on Animal Knowledge (CRCA), Middle for Integrative Biology (CBI); CNRS, Academy Paul Sabatier, Toulouse, France

Simon Klein, Cristian Pasquaretta, Jean-Marc Devaud & Mathieu Lihoreau
Department of Biological Sciences, Macquarie University, NSW, Australia

Simon Klein & Andrew B. Barron
Honeybee Research Institute, Jiangxi Agronomical University, Nanchang, Jiangxi, 330045, P.R. China

Xu Jiang He
Department of Biological and Experimental Psychology, School of Biological and Chemic Sciences, Queen Mary University of London, London, E1 4NS, UK

Clint Perry
Volda Academy College, Department of Science and Mathematics, Volda, 6100, Norway

Eirik Søvik

Contributions

Due south.K., E.Southward., C.J.P., M.L. and A.B.B. conceived the report and designed the experiment. S.Thou. and 10.J.H. conducted the experiments. S.Chiliad., C.P., Thousand.L. and A.B.B. analysed the information. S.Chiliad., C.P., X.J.H., C.J.P., Due east.Southward., J.M.D., Yard.L. and A.B.B. wrote the manuscript.

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Correspondence to Andrew B. Barron.

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Klein, S., Pasquaretta, C., He, X.J. et al. Love bees increment their foraging performance and frequency of pollen trips through feel. Sci Rep ix, 6778 (2019). https://doi.org/10.1038/s41598-019-42677-x

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Received: 21 August 2018
Accepted: 05 April 2019
Published: 01 May 2019
DOI : https://doi.org/x.1038/s41598-019-42677-10

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