Source: The University of North Carolina at Greensboro submitted to NRP
FINE-SCALE DYNAMICS OF HONEYBEE GUT MICROBIAL COMMUNITIES AFTER PERTURBATION
Sponsoring Institution
National Institute of Food and Agriculture
Project Status
COMPLETE
Funding Source
Reporting Frequency
Annual
Accession No.
1018799
Grant No.
2017-67012-29382
Cumulative Award Amt.
$62,401.32
Proposal No.
2018-02589
Multistate No.
(N/A)
Project Start Date
Apr 15, 2019
Project End Date
Sep 30, 2019
Grant Year
2019
Program Code
[A7201]- AFRI Post Doctoral Fellowships
Recipient Organization
The University of North Carolina at Greensboro
1400 Spring Garden Street
Greensboro,NC 27412
Performing Department
Biology
Non Technical Summary
Honeybees are responsible for pollinating crops that make up at least one-third of our diet, and their value to global agricultural production reaches many billions of dollars per year. Since 2006, the world's honeybee colonies have undergone elevated mortality, and annual colony losses continue to rise. The reasons for this phenomenon, commonly called colony collapse disorder, are not fully understood, but several possible causes have been suggested, such as pesticides, genetically modified crops, habitat fragmentation, climate change, and various diseases and parasites. However, no single stressor has been shown to account for the declines.Gut microbial communities influence animal health in many ways including synthesis of vitamins, balance of metabolism, defense against pathogens, behavior, and developmental and immune responses. Perturbed gut communities can be detrimental to host health. Honeybees are exposed to a number of chemicals, such as pesticides, herbicides, and antibiotics, which are potent sources of disturbance to microbial communities. In view of growing evidence for the importance of the gut microbial communities in animal health and the largely unexplained decline of honeybee colonies, the effects of chemical exposure on the honeybee gut microbiome are of major interest.The work proposed here will provide a full description of how exposure to pesticides, herbicides, and antibiotics affects the native gut microbial communities of honeybees. This project will also address how disruption of the healthy gut community affects susceptibility to invasion by opportunistic pathogens. These findings will provide insights on the importance of resident gut microbes in honeybee health and will be particularly relevant to beekeepers by revealing what types of perturbations of gut communities increase the risk for pathogen infection. Thus, the results from this project will provide insight into the decline of honeybee populations.
Animal Health Component
20%
Research Effort Categories
Basic
80%
Applied
20%
Developmental
(N/A)
Classification

Knowledge Area (KA)Subject of Investigation (SOI)Field of Science (FOS)Percent
3143010110050%
3153010110050%
Goals / Objectives
The proposed research will investigate the effects of chemically induced perturbation on the native gut microbiota of honeybee. These perturbations can have negative impacts on bee health, for example increased susceptibility to pathogen infection. I will determine the effects of antibiotics, pesticides, and herbicides on the native gut microbial community structure, and will evaluate whether recovery of the microbiota can be achieved after perturbation. Furthermore, it will address how disruption of the healthy community affects susceptibility to invasion by non-resident species, particularly opportunistic pathogens. To our knowledge, this is the first study to look at the effects of antibiotics, pesticides, and herbicides on the native honeybee gut microbial community structure.The long-term goal of this proposal is to understand the effects of perturbation on host- associated microbial communities. The honeybee, an agriculturally and economically important pollinator, will be used as a model system to address the following supporting objectives:Identify the impact of perturbation on the population dynamics of the gut microbiomeDetermine the ability of gut microbial community to recover after perturbationDetermine how perturbation of the gut microbiome promotes invasion by opportunistic pathogens.Findings of this project will provide insights into the importance of resident gut microbes for honeybee health.
Project Methods
Key to the proposed work is an experimental approach that will enable identifying fine-scale community changes that cannot be detected using 16S rRNA sequence profiling. As opposed to the highly conserved 16S rRNA, protein-coding genes evolve much faster. For example, bacterial strains differing by only 1% in 16S sequence can differ by 10-20% in single copy, universal protein-coding genes. This is directly observed in the bee bacteria. Thus, an approach based on protein-coding genes can reveal patterns of strain diversity and fine-scale differences in the community structure. This approach echoes an older method known as multilocus sequence typing (MLST), which is used to classify strains within species. Here, the critical difference is that it will be applied to obtain comprehensive characterizations of entire uncultured communities, and will exploit high- throughput sequencing technologies. Using the complete genomes from the core species of the bee gut microbiome, I have defined marker genes for each species. These markers were selected based on their universal presence in strains and their ability to capture maximal diversity. Using these markers I will be able to track strain level diversity much more accurately than is possible with 16S rRNA profiling. Moreover, traditional rRNA 16S profiling will also be applied in order to define the overall composition of the community (i.e. the presence of the major phyla).Objective 1. Identify the impact of perturbation on population dynamics of the gut microbiomeThe Raymann lab maintains five hives on the UNCG North campus and has established detailed protocols for experimentally manipulating gut microbiota of honeybees. NEWs will be collected from a removable frame kept overnight in an incubator. After emerging, the bees will be marked and released back into the colony so they can naturally acquire their microbiota. After one week, I will collect marked bees and subject them to i) antibiotics (tylosin) ii) imidacloprid iii) glyphosate and iv) combinations of the three treatments. Controls will be fed sterile sugar syrup. Cohorts of bees will be dissected before and after chemical treatment. DNA will be extracted from the gut using established techniques, and Illumina-based amplicon profiling of 16S rRNA and defined protein-coding gene markers will be performed. I will also estimate the total community size pre- and post-treatment by quantifying 16S rDNA copies using qPCR (adjusting for number of rRNA operons per genome). All DNA sequence data will be analyzed using bioinformatic approaches including sequence comparison, estimation of gene polymorphism, identification of operational taxonomic units (OTUs), and phylogenetic and taxon-based analysis of diversity within and between samples. Pilot experiments based on Illumina sequencing of the V4 region of the 16S rRNA gene have confirmed that treatment with tetracycline has a significant impact on the community structure of the microbiota.Objective 2: Determine the ability of the gut microbial community to recover after perturbationI will perform controlled experiments to test if different environmental factors play a role in the recovery of the gut microbial community after chemical perturbation (described in Objective 1). These experiments will involve allowing the bees to recover after chemical treatment in solitary cup cages, in cup cages containing normal, untreated workers taken from the hive, and in intact hives outdoors. The recovery groups maintained in cup cages will be fed with a sterile diet of sugar syrup. The bees will be allowed to recover for two weeks, and survival rates will be recorded daily. Cohorts of bees will be sampled at several time points during the experiment, and Illumina-based amplicon profiling will be done as described above.Objective 3. Determine how perturbation of the gut community promotes invasion by opportunistic pathogensSeveral opportunistic pathogens of the honeybee have been isolated in the lab and can be easily cultured. Newly emerged workers will be marked and released into the hive so they can naturally acquire their microbiota. After one week, I will collect marked bees and subject them to different chemical treatments (as described above). I will then expose the bees to various opportunistic pathogens of bees, e.g. Serratia marcescens, Escherichia coli, and Hafnia alvei. The colonization of the pathogen in the gut will be determined by plating the guts on LB agar plates (native gut members will not grow on this medium). Bees will be sampled (dissected) and survival rates will be monitored at multiple time points throughout treatment. DNA will be extracted from the gut using established techniques, and Illumina-based amplicon profiling of 16S rRNA and defined protein-coding gene markers will be performed to determine pre- and post-infection community composition. The relative abundance of the opportunistic pathogen will be compared to the entire community using qPCR. All DNA sequence data will be analyzed using the bioinformatics approaches described above.Progress towards completion of the project objectives will be based on each aim in the proposed timeline (2 years), which includes completing treatment, recovery, and infection experiments, data collection and analysis, and preparing manuscripts.

Progress 04/15/19 to 10/14/19

Outputs
Target Audience:Microbiologist, Environmental scientists, Microbiome researchers, beekeepers, farmers, veterinarians Changes/Problems: Nothing Reported What opportunities for training and professional development has the project provided?As a postdoctoral researcher this project allowed me to speak at several national and international conferences, publish several peer-reviewed research articles, and learn new experimental techniques. After obtaining a position as an Assistant professor this project allowed me mentor graduate and undergraduate students. How have the results been disseminated to communities of interest?Through prsentation at scientific conferences, publications, particpation in outreach events, news stories, and my personal website. What do you plan to do during the next reporting period to accomplish the goals? Nothing Reported

Impacts
What was accomplished under these goals? Effects of tetracycline on the gut microbiome and survivorship of honeybees My first project evaluated how the antibiotic tetracycline impacts the gut microbiota and survivorship of honeybees using 16S rDNA gene profiling, quantitative PCR, and survival assays. From this study I was able to show that tetracycline severely alters the gut community composition and size and leads to decreased survivorship in the hive and the lab. I found that the effects of tetracycline on the gut community were still very evident even one week after treatment was stopped, suggesting that bees do recover from tetracycline exposure. Furthermore, I found that treatment with tetracycline caused bees to be more susceptible to the pathogen Serraita marcesences, leading to over 50% mortality rate in treated bees. These results were published in PLoS Biology in 2017 and have received a great deal of media attention. Additionally, I have presented this work at one international conference and one regional meeting. Effects of tetracycline on the fine-scale diversity of the honeybee gut microbiome In a second project I investigated how tetracycline treatment impacts fine-scale diversity in the microbiome using species-specific protein-coding genes. This approach allowed me to follow two of the dominant core gut species over time following tetracycline treatment. In this study I found that the genetic diversity of the two core species investigated was drastically decreased following treatment. However, these two species were not equally impacted by treatment, suggesting that some community members are more resistant to tetracycline. My results show that this method provides the resolution necessary to follow fine-scale dynamics in natural microbial communities over time, something that could not be achieved by 16S rDNA gene profiling. The results from this study are published in Molecular Ecology. Effects of pesticides on the gut microbiome and survivorship of honeybees We have investigated how two different pesticides, glyphosate and imidacloprid impact the gut microbiome of honeybees. We found that glyphosate alters the bee gut microbiome composition which can lead to increased susceptibility to pathogen infection. We did not find any evidence that imidacloprid alters the gut microbiome. However, imidacloprid did reduce honeybee survival. These results have been published in two separate publications (Proceedings of the National Academy of Sciences and Applied Environmental Microbiology). Characterization of the honeybee pathogen S. marcescens We have characterized the virulence of the pathogen S. marcescens to honeybees. To do this three pathogenic S. marcescens strains were isolated from honeybee guts sequenced. These three strains were the first S. marcescens species to be characterized from the guts of honeybees. Through a series of experiments, we found that all three strains can be highly virulent to honeybees. These results have been published in MBio.

Publications

  • Type: Journal Articles Status: Published Year Published: 2018 Citation: Antibiotic exposure perturbs the gut microbiota and elevates mortality in honeybees K Raymann, Z Shaffer, NA Moran PLoS biology 15 (3), e2001861
  • Type: Journal Articles Status: Published Year Published: 2018 Citation: Antibiotics reduce genetic diversity of core species in the honeybee gut microbiome K Raymann, LM Bobay, NA Moran Molecular ecology 27 (8), 2057-2066
  • Type: Journal Articles Status: Published Year Published: 2018 Citation: The role of the gut microbiome in health and disease of adult honey bee workers K Raymann, NA Moran Current opinion in insect science 26, 97-104
  • Type: Journal Articles Status: Published Year Published: 2018 Citation: Imidacloprid decreases honey bee survival rates but does not affect the gut microbiome K Raymann, EVS Motta, C Girard, IM Riddington, JA Dinser, NA Moran Appl. Environ. Microbiol. 84 (13), e00545-18
  • Type: Journal Articles Status: Published Year Published: 2018 Citation: Glyphosate perturbs the gut microbiota of honey bees EVS Motta, K Raymann, NA Moran Proceedings of the National Academy of Sciences 115 (41), 10305-10310
  • Type: Journal Articles Status: Published Year Published: 2018 Citation: Pathogenicity of Serratia marcescens Strains in Honey Bees K Raymann, KL Coon, Z Shaffer, S Salisbury, NA Moran mBio 9 (5), e01649-18
  • Type: Journal Articles Status: Published Year Published: 2019 Citation: Population Genetics of Host-Associated Microbiomes LM Bobay, K Raymann Current Molecular Biology Reports 5 (3), 128-139
  • Type: Journal Articles Status: Under Review Year Published: 2019 Citation: Strain structure and dynamics revealed by targeted deep-sequencing of the honeybee gut microbiome LM Bobay, E Wissel, K Raymann Molecular Ecology
  • Type: Conference Papers and Presentations Status: Other Year Published: 2019 Citation: The microbiome of honey bees Virginia Tech Department of Entomology Seminar Series Blacksburg, Virginia
  • Type: Conference Papers and Presentations Status: Other Year Published: 2018 Citation: The microbiome and honey bee health Microbiome symposium. The Center for Root and Rhizobiome Innovation. Lincoln, Nebraska
  • Type: Conference Papers and Presentations Status: Other Year Published: 2018 Citation: Antibiotics and honey bees Presidential Advisory Council on Combating Antibiotic Resistant Bacteria public meeting Columbus, Ohio
  • Type: Conference Papers and Presentations Status: Other Year Published: 2018 Citation: Impact of antibiotics on honey bee health American Veterinary Medical Association National Convention Denver, Colorado
  • Type: Conference Papers and Presentations Status: Other Year Published: 2019 Citation: Antibiotics and honey bees Honey bee Veterinary Consortium Conferencce Raleigh, North Carolina
  • Type: Conference Papers and Presentations Status: Other Year Published: 2019 Citation: Community dynamics and evolution of the honey bee gut microbiome Microbiome Seminar Series at North Carolina State University Raleigh, North Carolina