Duration: 10/01/2009 to 09/30/2014

Administrative Advisor(s):

NIFA Reps:

Non-Technical Summary

Statement of Issues and Justification

ISSUES AND JUSTIFICATION

Honey bees provide essential pollination services to US fruit and vegetable growers, adding $8-10 billion annually to farm income. About 2 million colonies are rented by growers each year to service over 50 crops. Almonds alone require 1.3 million colonies and are predicted to require 2.12 million by 2012, a number nearly equal to the number of colonies presently in the US. Increasing demand comes at a time when beekeepers are operating in crisis mode. The supply of healthy colonies is volatile as parasitic mites and the rigors of migratory beekeeping continue to cause catastrophic die-offs. Pesticide resistance and a lack of viable, industry-based honey bee breeding programs contribute to these losses. The problems associated with mites and other factors affecting honey bees are outlined in a 2007 report of the NAS-NRC², Status of Pollinators in North America.

The winter of 2007/08 witnessed another major colony die-off, and while many of the deaths are due to parasitic mites, a large number of colonies exhibited symptoms inconsistent with mites or any known disorder. By last count, 0.75 to 1 million honey bee colonies died over the winter of 2007-2008 (vanEngelsdorp et al. 2008). Migratory beekeepers trucking bees over great distances were especially hard hit. This suggests yet another problem has beset an already beleaguered industry. This new syndrome has been named Colony Collapse Disorder (CCD) in 2007. A list of possible causes for CCD includes new pesticides and pesticide use patterns, nutritional deficits associated with monocultures, loss of immunity to pathogens and exotic pathogens.

According to a survey conducted by the Apiary Inspectors of America (AIA) and the U.S. Department of Agriculture, honey bee colony losses nationwide were approximately 29 percent from all causes from September 2008 to April 2009 (Kaplan 2009). This is less than the overall losses of about 36 percent from 2007 to 2008, and about 32 percent from 2006 to 2007, that have been reported in similar surveys. About 26 percent of apiaries surveyed in the latest survey reported that some of their colonies died of colony collapse disorder (CCD), down from 36 percent of apiaries in 2007 2008. As this was an interview based survey, it was not possible to differentiate between verifiable cases of CCD and colonies lost as the result of other causes that share the "absence of dead bees" as a symptom. However, among beekeepers that reported any colonies collapsing without the presence of dead bees, each lost an average of 32 percent of their colonies in 2008 2009, while apiaries that did not lose any bees with symptoms of CCD each lost an average of 26 percent of their colonies. The survey checked on about 20 percent of the country's 2.3 million colonies.

The NC 508 committee met at the University of Florida, Gainesville on 4 February 2009. Members present represented 14 universities from throughout the US. Committee members discussed ongoing research and plans were made to conduct future research to develop viable solutions to the problems afflicting honey bees in order to ensure the sustainability of the nations food supply.

Most of our committee members are involved in a $4.1 million 4-year CAP project that was funded in 2008 to study the causes of CCD and other maladies affecting bee populations. The CAP funding obtained was a direct result of the establishment of the NC 508 committee in 2007. A second meeting of some NC 508 participants was convened that same year to initiate the proposal writing. The scientists will conduct research that addresses genomics, breeding, pathology, immunology, and applied ecology to investigate and explain the causes of the decline in bee colonies in the US in recent years. In addition, we will investigate the role of ecosystem services provided by native, wild bees in pollinating crops. Native bee pollination can be sufficient to fully pollinate crops in some agricultural contexts (Kremen et al. 2002; Winfree et al. 2007), and even when pollination services are incomplete, can serve as a supplement to or back-up for managed honey bee stocks. Native bee ecologist(s) will investigate the role of native bee species in pollinating several crops and will and identify the land and farm management practices associated with high levels of native bee pollination. Committee members will work closely with the extension community and other stakeholders to develop and implement mitigative strategies that unravel the causes of CCD and other significant bee health problems.

TYPES OF ACTIVITIES

The purpose of this committee has been and will be to coordinate research that is relevant to bee colony health. We are seeking participation of personnel with expertise in bee nutrition, toxicology, parasitology, pathology, breeding, integrated pest management, and non-Apis species. Research and extension personnel will meet annually to discuss coordination and will form subgroups that will coordinate or collaborate on research throughout the year. Extension personnel will coordinate in technology transfer and adoption of research findings to beekeepers.

Related, Current and Previous Work

Most members of the NC508 committee are active in the American Association of Professional Apiculturalists (AAPA) and have been involved in research and extension that pertains to honey bee health. The NC 508 committee met at the Double Tree Hotel, Sacramento, California on 10 January 2008. Members present represented 14 universities from throughout the US. Committee members discussed ongoing research and plans were made to conduct future research to develop viable solutions to the problems afflicting honey bees in order to ensure the sustainability of the nations food supply.

Most of our committee members are involved in a $4.1 million 4-year CAP project that was funded in 2008 to study the causes of CCD and other maladies affecting bee populations. The CAP funding obtained was a direct result of the establishment of the NC 508 committee in 2007. A second meeting of some NC 508 participants was convened that same year to initiate the proposal writing. The scientists will conduct research that addresses genomics, breeding, pathology, immunology, and applied ecology to investigate and explain the causes of the decline in bee colonies in the US in recent years. Committee members will work closely with the extension community and other stakeholders to develop and implement mitigative strategies that unravel the causes of CCD and other significant bee health problems.

Objectives

1. To develop and recommend to beekeepers "best practices" for varroa mite control based on currently available methods and strategies for mite management.
   
2. To evaluate the role and causitive mechanisms of parasitic mites and pathogens such as viruses, protozoa and bacteria in honey bee colony deaths.
   
3. To determine the effects of pesticides and other environmental chemicals on honey bee colony health.
   
4. To determine how environmental factors, including nutrition and management practices affect honey bee colony health.
   
5. To determine the effects of interactions among various factors affecting honey bee colony health.
   
6. To coordinate research and extension efforts related to bee colony health.</li>
   
7. To facilitate, through research and extension activities, the development of industry-based honey bee stock selection, maintenance and production programs that demonstrably incorporate traits that confer resistance to pests, parasites and pathogens.
   
8. To focus on non-Apis bees, their conservation, pathology, susceptibility to pesticides, and their contribution to crop pollination including economic value.
   

Methods

1) The CAP program provides the fundamental framework for this effort. The team is made up of several research teams comprehensively addressing causes of bee decline, whether with pathogens or toxins, and feeding this information directly to the Bee Health Community of Practice website at eXtension.org. The public side of this site can be viewed at http://preview.extension.org/bee%20health. Community work pages have been populated with organizational information, website content outline, and 23 pages for the public site. Additional content is being obtained and is in preparation for upload, including bulletins in bee biology, best management practices, and diseases/pests. An online meeting protocol was established and we conducted meetings with the CoP, which included planning for the CAP team. This effort is being managed by a dedicated CAP-funded staffer, Mr. Michael Wilson at University of Tennessee. Collaborations: The Bee Health Community of Practice is so far comprised of 43 members representing scientists and extension workers from CAP, the ARS Areawide project, and numerous non-affiliated experts across North America. 2) Background: This huge effort is a shared goal of members of CAP as well as ARS Areawide and includes a significant fraction of the honey bee scientists of the United States. As this NC 508 project was the synergist for the CAP team, we will limit our reports in this section to objectives of this particular group. Researchers will coordinate their efforts geographically and methodologically in sampling for detecting parasites and pathogens in hives and bees. Pathogen identification in bees will be aided by molecular techniques (e.g. massively parallel DNA sequencing, bioinformatics and PCR). Causative relationships will be evaluated in part by correlating the presence of various biotic agents and colony symptoms over time. Objective 2a - Comparative virulence of Nosema apis and N. ceranae in caged bees. Newly emerged workers will be individually inoculated with 0 (control), 5, 50, 500, 5,000, and 50,000 spores with both species of Nosema. One hundred bees per cage (cage size: 14 × 12 × 16 cm), 4 cages per dosage will be used for each colony. Workers are provided with 50% sugar syrup (changed every 3 days), bee pollen (in which Nosema spores have been inactivated by heating at 60° C for 24 hrs), and distilled water ad libitum. Dead bees will be removed daily and their numbers recorded. The experiment will be repeated with bees from at least three different colonies. Survival analysis will be used to determine the time required for 50% mortality in the three doses of Nosema, for both species. The median infective dosage (ID50) will be calculated using probit analysis (Finney, 1971) for both species of Nosema. (For additional background, rational and significance,sub-objectives, expected outcomes, methods, and collaborators, see full/separate document) 3) Synergism bioassays - The LD50 for 8 miticides (Apistan, CheckMite, Mite Away II, Apiguard, Api Life Var, Mite-A-Thol, Hivastan, and Sucrocide) used in bee hives will be established using 360 worker bees per bioassay. All bioassays will be done by topically applying serial dilutions of compounds with a Hamilton micro syringe. All compounds will be tested singly and in reciprocal tests for all synergism bioassays. Synergistic ratios will be determined by treating bees with LD5 of one compound and then treating with serial dilutions of the second compound to determine whether the dose response curve of second compound shifts position. Synergistic ratios will be determined if synergism is observed. EPA guidelines for the age and diet of bees used in bioassays will be followed. Sperm number and viability and queen survivorship - The LD50 for newly emerged drones will be determined for each of the 8 compounds. Drones will be treated with non-treated controls and a sub-lethal dose as described above. Newly emerged drones will be marked with enamel paint upon emergence. They will be treated, released back into the colony, and then recaptured 14 days later for examination. More drones will be marked and treated than needed for the analysis, so that if some are lost on mating flights enough will remain for the assay. Marked drones will be captured 14 days later and their sperm counts and sperm viability determined by examining their sperm in a hemocytometer. Sperm will be collected from individual drones by applying pressure to the abdomen causing full eversion of the endophallus (Harbo 1985). The semen will be then collected using a capillary tube. Sperm viability will then be assessed using two fluorescent stains (Locke et al. 1990). Propidium iodide will stain the dead cells red and Hoechst No. 33342 (H342) will stain the living cells green. The stains will be added to sperm dilutions, and a drop of the stained solution will be placed on a microscope slide with a cover slip and examined at 400X magnification (Collins 2001). The number of dead (red) and live (green) cells in 2-3 samples per drone will be counted, and a percentage of live cells calculated. We will raise virgin queens for this assay. They will be emerged in caged cells in an incubator. They will be treated and placed in individual queen cages. All queens will be placed into holding frames and transferred to a queen bank. Queen bees will be treated with a dose of each of the 8 compounds that is sub-lethal for worker bees, including non-treated controls. Queens in the queen bank will be examined weekly for at least 30 days for mortality, and emerging brood will be added to the queen bank each week to keep it vigorous. For each compound test, 30 treated queens and 30 untreated queens will be maintained. Collaborators: Ellis, M. (For additional background, rational and significance,sub-objectives, expected outcomes, methods, and collaborators, see full/separate document) 4) Objective 4a  Determine whether there is an interaction between nutritional status and Nosema disease. Newly emerged workers will be individually inoculated with 10,000 spores on day 0 and provided with no pollen, mixed-bee pollen or monocultural pollen from a few representative plants important for honey bee pollination (cherry, almond, corn, etc). Bees are provided with 50% sugar syrup (changed every 3 days) and distilled water ad libitum. One hundred bees per cage (cage size: 14 × 12 × 16 cm), 4 cages per treatment will be used for each colony. Longevity of workers will be compared among the treatments by survival analysis using SAS 9.1.3. Collaborators: Huang Other Research: A survey of management practices will be conducted through an eXtension website as part of the honey bee CAP. (For additional background, rational and significance,sub-objectives, expected outcomes, methods, and collaborators, see full/separate document) 5) Objective 5a  Effect of pathogens and stresses on survivorship of life-stage and caste cohorts One hundred bees per cage (cage size: 14 × 12 × 16 cm), 4 cages per treatment from three colonies will be obtained (3500 bees or 35 cages = 4 cages per treatment X 3 treatments X 3 colonies). Subject to the treatment variables under investigation, workers are provided with 50% sugar syrup (changed every 3 days), bee pollen and distilled water ad libitum. Dead bees will be removed daily, their numbers recorded and analyzed for pathogens. The experiment will be repeated with bees from at least three different colonies selected based on pathogen profile. To test the impacts of poor nutrition on bees, cage studies will first be tested with a combination of pathogens, using data from above to guide the selection of time points and levels to test. Nutrition will be examined using comparison of honey/bee bread (sterilized via gamma radiation and checked for sterility and nutritional changes), and sugar syrup/artificial bee pollen. The later will be manipulated to obtain diets of different nutritive value for the bees. Newly emerged workers divided into 3 treatments (control with normal diet, optimal artificial diet, and suboptimal diet with known poor bee performance). Temperature will be examined through a design similar as described above. Caged, newly emerged bees in replicates from the same colony will be exposed to temperatures ranging from 15° C to 40° C. These studies will be conducted at Illinois and samples sent for molecular analysis to PSU. Collaborators: Cox-Foster, Ostiguy, Solter (For additional background, rational and significance,sub-objectives, expected outcomes, methods, and collaborators, see full/separate document) 6) Objective 6a  Establish a Managed Pollinator Community of Practice with eXtension and populate website with new literatures on Best Management Practices and Bee Conservation The project budgets for a dedicated technician to work under J. Skinner to execute our eXtension initiatives. We will initiate and maintain a Community of Practice (CoP) on Managed Pollinators which will serve interested clienteles  the Community of Interest or CoI. The CoP will be the main conduit for media-based deliverables streaming from this CAP. Co-investigators will direct new information to the eXtension technician who will prepare Best Management and Conservations literatures and recommendations to accompany diagnostic reports. We will develop and share this CoP in full collaboration with ARS scientists involved in the Areawide project (J. Pettis, ARS Areawide Coordinator, see Documentation of Collaboration). Collaborators: Skinner (For additional background, rational and significance,sub-objectives, expected outcomes, methods, and collaborators, see full/separate document) 7) Objective 7a  Identify genes that confer resistance to Varroa and pathogens, and genes that respond to biotic challenges Gene responses to infection - Two control colonies and two colonies infected with N. ceranae will be established. A second set of experiments will use colonies infected with IAPV. Nurse bees will be collected from brood frames onto dry ice and screened for Nosema or viral infection. RNA will be extracted from the immune tissues (abdominal fat bodies) of pools of 5 bees of known age. Gene expression of infected and control bees will be compared on whole genome microarrays as described below, with six biological replicates/colony. Gene expression differences will be analyzed using ANOVA. Genes that are significantly different between healthy and infected colonies will be analyzed to determine if any specific functional categories are affected (Gene Ontology analysis). For example, we should be able to determine if specific genes within pathways related to immune response are affected (Aronstein and Salvidar 2005; Evans at al. 2006). Once candidate genes have been identified, qRT-PCR will be used to monitor expression of these genes in a broader range of treatment groups (Aronstein et al. 2006). Mapping disease and parasite resistance genes - QTL influencing VSH will be identified in one backcross family after analyzing a panel of 1,536 SNPs in 192 worker bees using the Illumina BEAD station. We will make a small hole in newly sealed worker pupal cells and infest them with individual Varroa mites. We will sample at least 96 bees on dry ice that are observed opening these cells, and 96 same-age control bees from the same backcross family. For each SNP, genotypes of bees performing the behavior will be compared with control bees in 2X2 Chi-Square tests to identify loci that have significantly different frequencies of marker-alleles in bees performing the behavior (Arechavaleta-Velasco and Hunt 2004). Prior to SNP genotyping, DNA from the drone father of the F1 queen and the F1 queen herself will be subjected to sequencing on an ABI SOLiD sequencer for SNP discovery. Sequence reads will be aligned with genomic contigs and SNPs identified using PolyBayes software, targeting SNPs within or near coding exons to associate specific genes with the trait. Probes for SNP genotyping will be designed. Similar techniques utilizing a backcross family segregating for disease resistance will identify QTL and candidate genes that influence resistance. The quantitative trait measured in this case will be the pathogen levels within bees as determined by qRT-PCR. Mapping eQTL involved in honey bee resistance mechanisms - Genomic DNA will be isolated from abdomens from the backcross family used to map QTL for VSH and from heads and thoraces of bees used to map resistance to pathogens (see above). This will leave heads to be analyzed for gene expression associated with behavior and abdomens containing fat bodies for expression of genes involved in immune responses. cDNA will be obtained from these same individuals, which will then be transcribed into labeled RNA and hybridized to a microarray containing nearly all of the annotated genes in the honey bee genome (Grozinger et al. 2007). eQTL will then be analyzed in the VSH mapping population and the pathogen-resistance QTL population. Crosses to map QTL for VSH and microarrays in this population will be conducted by the Baton Rouge bee lab with separate funding (see Danka, Documentation of Collaboration). Collaborators: Hunt, Spivak, Webster, Aronstein, Grozinger (For addtional background, rational and significance,sub-objectives, expected outcomes, methods, and collaborators, see full/separate document) 8) Objective 8a  Identify and characterize pathogens of managed non-Apis bees and determine cross-infectivity of microbes and parasites with Apis Commercially-produced and wild non-Apis bees will be surveyed for microbes (microsporidia, including Nosema; viruses, bacteria, and trypanosomes). Metagenomic methods developed for Apis (CAPS Project) will target microbes present in Koppert Bombus impatiens, US- and Canadian-produced Megachile rotundata, and wild foraging Bombus spp., Osmia spp., and Megachile spp. collected at five geographical locations (Maine, Massachusetts, Florida, Tennessee, New York) in the eastern United States. For commercial Bombus, 10 research colonies will be deployed at crop sites at each state; prior to deployment, 5-10 individuals will be removed and frozen at -20° C until processing. After 6 weeks of foraging in contact with wild bees, all adults will be recaptured and frozen for studies to determine development of pathogen load and presence of new microbe species. For commercial Megachile, 200 adults emerging from each of four vendors will be analyzed. Wild bees will be collected, identified to species, and pooled by species and location for analysis. Prior to processing, each wild bee will be inspected for parasites and assessed for pollen load and age. Information on the wild Bombus species pathogens will augment data produced for Bombus species in the western and Midwestern states by a current NRI grant to Cameron/Solter/Strange/Griswald. These researchers are working on genetic tests to distinguish N. bombi from other microsporidian species. Collaborators: Averill and participants in stationary apiaries, objective 3d) Objective 8b  Elucidate lethal and sub-lethal effects of insecticides on non-Apis General rationale and significance Many insecticides are toxic to bumble bees, particularly after direct spray or through exposure to treated foliage (Thompson 2001). However, there is a void in toxicity tables for non-Apis, particularly for newer chemistries. Pesticide toxicities determined for honey bees are not always predictive of toxicities to other bees NRC (2007). There is debate on the extent to which neonicotinoids accumulate in pollen and nectar and impact bee health. In the case of non-Apis bees, the results are mixed, with some showing no effect (Franklin et al. 2004), and others showing sub-lethal impacts on Bombus activity (Gels et al. 2002), behavior, and learning (Morandin and Winston 2003). There is need to expand this knowledge base with the most commonly used insecticides and most commonly cultured non-Apis bees. Expected outcomes To gain: (1) New information on basic toxicology of some of our most common non-Apis managed pollinators and (2) new information on sub-lethal effects of neonicotinoids on non-Apis bees. (For addtional background, rational and significance,sub-objectives, expected outcomes, methods, and collaborators, see full/separate document)

Measurement of Progress and Results

Outputs

• Published best management extension manual for IPM of parasitic mites
• Comprehensive educational website on Bee Health on eXtension.org in cooperation with the
• Surveys of incidence and coincidence of pathogens and parasites of honey bees.
• Surveys of environmental contaminants in the field and in colonies and correlations with colony health..
• Studies of effects of sub-lethal doses of environmental contaminants on bees at ambient concentrations.
• 6. Reports on breeding programs that reduce parasite and pathogen impact. 7. Recommendations on the efficacy of non-Apis pollinators for several crops and agricultural settings, and recommendations on non-Apis pollinator conservation. 8. Reports of annual meetings. 9. Reports on breeding programs that reduce parasite and pathogen impact.

Outcomes or Projected Impacts

• Increased beekeeper awareness and adoption of best practices to maintain colony health by controlling Varroa mites with minimum use of pesticides.
• Increased knowledge of pathogen identities and levels within bee hives.
• Increased understanding of causative agents involved in the colony collapses.
• Demonstrable progress towards developing more resistance to Varroa mites and pathogens in breeding stocks of bees.
• Leverage of increased funding for research of colony health issues.
• (For additional expected outcomes and impacts, see full/sepatate document)

Milestones

(2009): Website on bee health on eXtension.org established and populated with bulletin on bee biology Best management practices draft #1 completed. Progress report on all objectives due at annual meeting in January 2010. First year of studies involving cage inoculations and monitoring of pathogens in colonies. Apiaries will be established in six geographic regions. First analyses to identify relatively resistant and susceptible lines for at least one pathogen. Assessment of virulence of pathogens and impact of pesticides. Establishment of eXtension community of practice. Collection of samples to assess genetic diversity in bees.

(2010): Research report draft #1 on effect of sub-lethal doses of environmental contaminants. Survey of environmental contaminants in the field and in colonies and correlations with colony health complete. Initial report on recommendations on non-Apis pollinator conservation. Progress report on all objectives due at annual meeting in January 2011. Conducting first crosses to map genes influencing resistance to mites. Successful implementation of eXtension website. Publication of Best Practices guide.

(2011): Complete and publish final research report on effect of sub-lethal doses of environmental contaminants. Initial breeding program report due. Initial report on efficacy of non-Apis pollinators for several crops and agricultural settings. Progress report on all objectives due at annual meeting in January 2012.

(2012): Final report of survey of incidence and coincidence of pathogens and parasites of honey bees and publish on eXtesnion.org. Initial report on the effects of interactions among various factors affecting honey bee colony health. Progress report on all objectives due at annual meeting in January 2013 and plans for submitting new project for next 5-year project (2015-2019).

(2013): Breeding program final report due and published on eXtension.org. Final report on efficacy of non-Apis pollinators for several crops and agricultural settings and publish on eXtension.org. Final report on the effects of interactions among various factors. affecting honey bee colony health and publish on eXtension.org. Final report on recommendations on non-Apis pollinator conservation and publish on eXtension.org. Final report on all objectives due at annual meeting in January 2014. Submit new project proposal for next 5-year cycle.

Projected Participation

View Appendix E: Participation

Outreach Plan

Results of the project will be made available on our eXtension website. We have developed a community of practice and are gathering content for the site. This will provide quick turnaround between research trials and extension delivery.

Members of the project will also conduct workshops with beekeepers and speak at beekeeper meetings. We are considering publishing a bimonthly progress report on various aspects of the project in the two biggest apicultural trade journals.

Organization/Governance

We propose to have a chair and a secretary. The secretary is responsible for meeting minutes and annual reports. The chair is responsible for planning and running the annual meeting and coordinating proposal writing.

The secretary is elected at the annual meeting and becomes chair in the following year.

Literature Cited

Citations:

¹2008 CSREES Managed Pollinator Coordinated Agricultural Project

²2007 report of the NAS-NRC, Status of Pollinators in North America.


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