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Survey of Corylus Resistance to Anisogramma Anomala From HORTSCIENCE 45(5):832–836. 2010. the ‘Gasaway’ source of resistance has been widely used in the OSU hazelnut breeding program, leading to the release of EFB- Survey of Corylus Resistance resistant pollinizer cultivars (Mehlenbacher and Thompson, 1991b; Mehlenbacher and to Anisogramma anomala from Smith, 2004) and cultivars with kernel qual- ity suitable for commercial production Different Geographic Locations (Mehlenbacher et al., 2007, 2009). Although the ‘Gasaway’ allele continues Thomas J. Molnar1, Joseph C. Goffreda, and C. Reed Funk to provide a high level of EFB resistance in Department of Plant Biology and Pathology, Foran Hall, 59 Dudley Road, the Pacific Northwest (PNW), breeders and School of Environmental and Biological Sciences, Rutgers University, New plant pathologists are concerned with the long-term durability of using only one source Brunswick, NJ 08901-8520 of single-gene resistance (Osterbauer, 1996; Additional index words. hazelnut, filbert, eastern filbert blight, Corylus avellana, disease Coyne et al., 1998; Pinkerton et al., 1998; resistance, nut breeding Lunde et al., 2006). Adding to this concern is the question whether the genetic diversity of Abstract. Anisogramma anomala (Peck) E. Mu¨ller is the causal agent of the disease A. anomala found in the PNW is less than eastern filbert blight (EFB) of hazelnuts (Corylus spp.). Little is known of its genetic that found across its native range because diversity and pathogenic variation. Most sources of host resistance have been identified in A. anomala in the PNW is believed to trace the Pacific Northwest, a region outside the native range of A. anomala believed to have back to a single point introduction in south- limited diversity of the fungus due to a long history of quarantine and its relatively recent west Washington (Gottwald and Cameron, inadvertent introduction. In an attempt to investigate the pathogenic variation of 1980; Johnson et al., 1996). Based on the A. anomala, 12 hazelnut genotypes that showed complete resistance in Oregon were pathogen’s wide native range and sexual inoculated with 12 isolates collected from across its native range. At the conclusion of reproduction, it is likely that genetic diversity the study, ‘Grand Traverse,’ ‘Ratoli’, OSU 541.147, OSU 495.072, and OSU 526.041 and pathogenic variation exists in the species. remained free of disease. ‘Closca Molla’, OSU 759.007, and OSU 587.044 were infected Consequently, A. anomala may exist outside by most isolates. ‘Gasaway’ was infected by the Michigan isolate, which was also the only the PNW with the ability to overcome ‘Gas- one to infect its offspring ‘Zimmerman’, although the lesion lacked sporulating stromata. away’ resistance. In an attempt to investigate Interestingly, ‘VR20–11’, another offspring of ‘Gasaway’, was infected by isolates from this question, Osterbauer (1996) subjected New Jersey, Minnesota, and Michigan. The Michigan isolate also caused the only signs of trees of VR6–28, an OSU selection contain- infection on OSU 408.040. ing the ‘Gasaway’ allele, to greenhouse in- oculations with A. anomala collected from across the eastern United States and Canada. Anisogramma anomala (Peck) E. Mu¨ller Turkey, which generally produces 60% to 70% At the conclusion of her experiment, none of is the incitant of the disease eastern filbert of the world’s crop (world total was 776,890 the isolates were able to incite typical EFB on blight (EFB), which causes severe cankering, tons in 2007). Turkey is followed by Italy, VR6–28. However, concern was raised when branch dieback, and the death of most Euro- which produces around 17% of the world’s isolates from Minnesota and Ontario caused pean hazelnuts, Corylus avellana L. It is an total, and then the United States, which pro- development of sunken non-sporulating le- obligate biotrophic pyrenomycete native to duces less than 5% (FAOStat, 2009). Ninety- sions. While these lesions did not form the a wide geographic area east of the Rocky nine percent of the United States hazelnut crop conspicuous ‘‘football-shaped’’ stromata typ- Mountains where it is found associated with is produced in the Willamette Valley of ical of the fungus, their presence suggested its much more tolerant natural host, Corylus Oregon (Mehlenbacher and Olsen, 1997). possible variability in the pathogen. Thus, americana Marshall (Fuller, 1908; Weschcke, The destructive nature of EFB was known Osterbauer’s findings reinforced the need to 1954; Farr et al., 1989; Johnson and Pinkerton, and quarantine laws were established in the maintain quarantine regulations in the PNW 2002). Anisogramma anomala is known to early 1900s to prevent its introduction into and to continue the search for additional reproduce only by ascospores and has a multi- the western United States (Barss, 1921; sources of genetic resistance to A. anomala. year lifecycle that requires the host plant to Lagerstedt, 1979). Despite these precautions, Lunde et al. (2006) investigated the cycle through a dormancy period after in- EFB was discovered in a commercial orchard inheritance of EFB resistance in progeny fection to express disease symptoms (Stone in southwest Washington in the late 1960s derived from ‘Zimmerman’, which carries et al., 1992; Pinkerton et al., 1993). EFB is be- (Davison and Davidson, 1973). Since then, it the ‘Gasaway’ resistance allele (G¨okirmak lieved to be the primary reason commercial has spread southward throughout the entire et al., 2009). Along with reporting that hazelnut orchards were never successfully Willamette Valley of Oregon where it threat- ‘Zimmerman’ typically confers resistance to established in the eastern United States (Barss, ens the long-term viability of the U.S. hazelnut progeny in a 3 resistant:1 susceptible ratio, 1921; Thompson et al., 1996). Alternatively, industry (Mehlenbacher, 2005). EFB control which is different from the 1:1 ratio observed hazelnut production thrived in western Wash- measures have been developed, including fun- in progeny of ‘Gasaway’, they found several ington and Oregon due to being outside the gicide sprays and therapeutic pruning; how- seedlings that expressed small sunken lesions, native range of A. anomala, as well as having ever, they are expensive, yield-reducing, and but with no sporulation. These seedlings a climate well suited for European cultivars not entirely effective (Johnson et al., 1996; amplified the RAPD markers UBC152–800 (Thompson et al., 1996). Currently, the top Julian et al., 2008). Therefore, the development and UBC268–580 that are closely linked to hazelnut-producing country in the world is of cultivars with genetic resistance to the path- the ‘Gasaway’ allele (Mehlenbacher et al., ogen appears to be the most effective means 2004), signifying its presence. Their findings for control (Mehlenbacher, 1994). Breeders at suggest that although progeny may not be Oregon State University (OSU; Corvallis, OR) ‘‘completely resistant’’ or ‘‘immune’’ as previ- Received for publication 9 Dec. 2009. Accepted for have been working on this objective since ously reported, ‘Gasaway’ and ‘Zimmerman’ publication 2 Feb. 2010. 1976, when the first controlled pollinations still transmit a very high level of resistance to We would like to acknowledge the contributions of J. Capik and M. Bergen, and the support of the were made with the obsolete pollinizer ‘Gas- their offspring. In addition, their work sug- Rutgers Center for Turfgrass Science and the New away’. ‘Gasaway’ was shown to carry a dom- gests that further investigation is needed to Jersey Agricultural Experiment Station. inant allele at a single locus that confers elucidate the response of the ‘Gasaway’ allele 1To whom reprint requests should be addressed; complete resistance to EFB (Mehlenbacher when expressed in different genetic back- e-mail [email protected]. and Thompson, 1991a). Since its discovery, grounds. 832 HORTSCIENCE VOL. 45(5) MAY 2010 MISCELLANEOUS Since the discovery of ‘Gasaway’, addi- Materials and Methods potted in 3.7-L plastic containers using tional sources of resistance to A. anomala a peat-based planting medium (PromixÒ have been identified at OSU, including sev- Twelve hazelnut genotypes shown to BX; Premier Horticulture, Rivie`re-du-Loup, eral C. avellana cultivars and selections as express complete resistance to A. anomala Que´bec), top-dressed with 5 g of 5- to 6- well as other Corylus species and inter- in Oregon, comprised of six named cultivars month release fertilizer (Osmocote Plus specific hybrids (Coyne et al., 1998, 2000; and six OSU breeding selections (Table 1), 15N–9P2O5–12K2O with micronutrient; The Lunde et al., 2000; Chen et al., 2005, 2007; were inoculated with A. anomala isolates Scotts Co.), and were grown in a greenhouse Sathuvalli, 2007). However, these new sour- collected across the pathogen’s native range maintained at 24 °C day/18 °C night. After ces were identified by challenging them with (Table 2). Five susceptible cultivars were 7 to 8 weeks, actively growing plants were A. anomala originating in the PNW, and their included as controls (Table 1). In general, moved into humidity chambers for inocula- response to isolates originating across the three replications of each genotype were tions. Following inoculations, plants remained pathogen’s native range is largely unknown. challenged in 12 separate disease inoculation in the greenhouse until early August, were The objectives of this study were to develop treatments (six in 2003 and six in 2004) and then moved outside under shade for 3 weeks, a better understanding of A. anomala’s path- their responses were evaluated after cycling and then were planted in the field at the ogenic variation and to assess the broader through two periods of dormancy. Rutgers University Vegetable Research and response of available sources of host resis- Dormant rooted layers and scion wood Extension Farm (North Brunswick, NJ) for tance by challenging a diverse group of were provided by OSU or the USDA Agri- subsequent evaluations. cultivars and selections shown to be com- cultural Research Service National Clonal Each isolate consisted of populations of pletely resistant in Oregon with A.
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