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1. Cover Page MERC Grant No. TA-MOU-08-M29-076 Prevalence 1. Cover page MERC Grant No. TA-MOU-08-M29-076 Prevalence of honeybee viruses in the Middle East and their association with the appearance of Colony Collapse Disorder (CCD). FINAL REPORT Covering Period: 2009-2014 Submitting Institution: National Center for Agriculture Research and Extension, Bee Research department. P.O. Box 639-Baq'a 19381. Jordan. Principal Investigator: HADDAD, N., National Center for Agricultural Research and Extension, Bee Research Department. P.O. Box 639-Baq'a 19381. Jordan, Tel.: (06) 4725071, Fax (06) 4726099, E-mail: [email protected] Co-Principal Investigators: Sela, I., The Hebrew University of Jerusalem, Faculty of Agricultural, Food, and Environmental Quality Sciences Rehovot 76100, Israel. Tel: 00972-8-9489377, E-mail: [email protected]. Submitted to: U.S. Agency for International Development Bureau for Global Programs Field Support and Research Center for Economic Growth and Agricultural Development 2- Table of Contents 1. COVER PAGE .................................................................................................................................... 1 2. TABLE OF CONTENTS....................................... .................................................................... 3 . 3- EXECUTIVE SUMMARY: ................................................................................................................. 5 4 - RESEARCH OBJECTIVES: ............................................................................................................... 6 5 - METHODS AND RESULTS:.......................................................................................................... 1 6. IMPACT RELEVANCE AND TECHNOLOGY TRANSFER: ................................................... 34 7. PROJECT ACTIVITIES/OUTPUTS: ............................................................................................. 8 8. PROJECT PRODUCTIVITY: ........................................................................................................... 404 9. FUTURE WORK: .............................................................................................................................. 44 4 3- Executive Summary: Honeybees play an important role in nature and agriculture, where they are needed for the pollination. In recent years; there have been increasing reports of substantial losses of honeybee colonies in many countries. These mortalities have multiple causes, from the still mysterious rapid adult depopulation phenomena and acute poisoning by novel to the more common mortalities due to honeybee parasites and pathogens, such as Nosema, foulbrood and in particular the parasitic mite Varroa destructor and the lethal virus epidemics it initiates and vectors. In the Middle East North Africa Region (MENA), abnormal mortality and local disappearances of bees are often reported by beekeepers; however this project is the only project to our knowledge and according to the available scientific publications that worked cross borders from throughout the area from Morocco until Iraq. Originally this project was designed to work only in some of the Middle East and on honeybee viruses mainly the IAPV virus, but due to the outstanding accomplishments since early stages of the project, the work was expanded to cover most of the MENA region and several other honeybee diseases. The Arab-Israeli collaboration was very strong in this project and we have been able to work on capacity building of several experts from 11 Arab countries and a reference laboratory is found in Jordan for the identification of honeybee diseases and viruses. Many diseases where identified and recorded official for the first time in many countries. During the project we communicated with many beekeepers both in Israel and in Jordan, and raised the awareness for honeybee diseases. 4 - Research Objectives: 4.1 Overall objective:- The major aim of the project was to study the extent of honey be viruses presence in the Middle East. This was important since viruses and mainly the Israel Acute Paralysis Virus (IAPV) were closely associated with Colony Collapse Disorder (CCD), which is a major economic problem world-wide, and to study the relation between finding the IAPV segment in the bee genome to the CCD and other factors causing the bee mortality and loss. 4.2 - A survey of bee viruses in the Middle East: In recent years there have been increasing reports of substantial losses of honeybee colonies in many countries both with respect to increased winter mortality and population losses during the active season (vanEngelsdorp and Meixner 2010). These mortalities have multiple causes, from the still mysterious rapid adult depopulation known as Colony Collapse Discorder (CCD; vanEngelsdorp et al. 2009) and acute poisoning by novel insecticides (Rosenkranz) to the more common mortalities due to honeybee parasites and pathogens, such as Nosema (Fries 2010), foulbrood (Genersch 2010; Forsgren 2010) and particularly Varroa destructor (Rosenkranz et al. 2010) and the lethal virus epidemics it initiates and vectors (de Miranda & Genersch 2010; de Miranda et al., 2010). Many research studies and surveys were conducted on the various causes of these mortalities and pathogens (Desjardin and Boucher, 2005; Ellis and Munn, 2005; Oldroyd, 2007; Burgett et al., 2009, Currie et al., 2010 , Guzman-Novoa et al., 2010; Carreck and Neumann, 2010; Jabaji and Copley, 2012). In the Arab world, abnormal mortality and local disappearances of bees are often reported by beekeepers in the years 2007-2010 (Haddad, 2011), but only few research publications in the MENA region where published on this subject before they beginning of this project, by the end of this project the picture became clearer due to the published findings by the network of this project research team. The honeybee Apis mellifera is host for many viral infections most of which generally persist without obvious symptoms, and only occasional virulent outbreaks (Bailey et al., 1981; Bailey and Ball, 1991; de Miranda et al., 2012). About 18 different viruses have been detected in honeybees so far, most of which have a small single- stranded RNA genome of positive polarity. The relationship of a number of these viruses with elevated honeybee colony mortality is a world-wide concern (Benjaddou et al., 2001; Grabensteiner et al, 2000; Bakonyi et al, 2002; Tentcheva et al , 2004, Chen et al., 2005; Genersch and Yue, 2005; Chantawannakul et al, 2006; Chen and Siede, 2007; Weistein et al., 2008; Genersch and Aubert, 2010). The principal virus associated with elevated colony mortality is Deformed wing virus (DWV). The virus was first isolated in the 1970’s from a sample of asymptomatic dead honeybees from Egypt, and called Egypt bee virus (EBV), before it was found to be serologically closely related to a virus from DWV-symptomatic bees from Japan, isolated in the early 1980’s (Ball and Bailey, 1991). Deformed wing virus (DWV) is a positive-strand RNA virus that was originally isolated for the first time from adult bees (Apis mellifera) and Varroa destructor in Japan (Bailey and Ball, 1991). The main host of DWV is Apis mellifera, where it is present in the world (Allen and Ball, 1996; Nordstrom et al, 1999. Calderon et al, 2003. Tentcheva et al, 2004. Berenyi et al, 2006. Chantawannakul et al, 2006. Forgach et al, 2008. Sanpa and Chantawannakul, 2009). This is the virus is already widely studied in recent years (Grabensteiner et al, 2000.. Huang, 2000. Benjeddou et al, 2001; Bakonyi et al, 2002.. Tentcheva et al, 2004. Chen et al, 2005. Genersch and Yue, 2005; Chantawannukul et al, 2006. Williams et al, 2009;. Martin et al, 2010;. Zioni et al, 2011;. Yanez et al, 2012). DWV has become one of the major virus threats to the honey bee industry due to its synergism with V. destructor (Martin, 2001), which acts as an inducer and highly efficient vector of the virus (Ball, 1989; Bowen-Walker et al., 1999; Nordström, 2000; de Miranda and Genersch 2010). The mites cause immunosuppression in the bees, activating otherwise nonlethal DWV infections which are efficiently transmitted by the mite precipitating a lethal epidemic that increasingly also involves other transmission routes (oral-fecal and sexual-vertical; de Miranda and Genersch 2010). In addition, the viral infections increase susceptibility to other opportunistic pathogens (Yang and Cox-Foster, 2005), leading to a complex disease profile at colony level (Shimanuki et al., 1999; Tentcheva et al., 2004; de Miranda et al., 2012) and a progressive reduction in colony performance, followed by large-scale death. In the absence of mite DWV is largely innocuous and often even undetectable, such that effective mite control stops and even reverses the DWV epidemic (Rosenkranz et al., 2010). Typical symptoms exhibited by severely DWV-infected bees are crumpled wings, bloated abdomens, paralysis, learning impairment, and a drastically shortened life span. There is a direct correlation between the viral titer and the symptoms displayed (Bailey and Ball, 1991) and the virus is chiefly responsible for the symptoms (Möckel et al., 2011). DWV has been detected in all honey bee life stages, as well as glandular secretions (Chen et al., 2005; Yue and Genersch, 2005), through which it is orally transmitted to larvae. DWV is also infectious in bumble bees, where it causes similar symptoms (Genersch et al., 2006), and is an apiculture-derived threat to wild pollinators (Fürst et al., 2014; Singh et al., 2010; Evison et al., 2011). The complete viral genome was
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