Eastern filbert blight (EFB), caused by Anisogramma anomala, is a devastating disease of Corylus avellana, the European hazelnut of commerce, and is considered the primary limiting factor of production in eastern North America. Conversely, C. americana, the wild American hazelnut, is generally highly tolerant of EFB, although it lacks many horticultural attributes necessary for commercial nut production. Hybrids of C. americana and C. avellana combine the EFB resistance of the wild species with the improved nut quality of the European species. However, inheritance of EFB resistance from C. americana remains unclear with existing hybrids derived from a very limited selection of parents. To investigate this topic, C. americana and advanced-generation C. americana × C. avellana hybrids were crossed with susceptible C. avellana and the resulting seedlings exposed to EFB through field inoculations and natural disease spread. In the winter after their fifth growing season, plants were rated for the presence of EFB using an index of 0 (no disease) through 5 (all stems containing cankers). The three progeny related to C. americana ‘Rush’ segregated for resistance in a ratio of one resistant to one susceptible, suggesting the presence of a single dominant R gene. A wide array of disease responses was observed for the other progenies with some expressing little EFB resistance or tolerance and others showing a distribution of disease phenotypes typical of control by multiple genes. Overall, the results indicate that both qualitative and quantitative resistance is present in C. americana. They also suggest that the choice of C. americana parent as well as the C. avellana parent will play a significant role in obtaining useful levels of EFB resistance in hybrid offspring, although the degree of disease expression in the parents may not be a useful predictor of progeny performance. Thus, more research is needed to understand inheritance of resistance, especially in advanced-generation backcrosses to susceptible C. avellana.
Thomas J. Molnar and John M. Capik
John M. Capik and Thomas J. Molnar
One hundred ninety clonal accessions of Corylus, including species and various interspecific hybrids of C. avellana, C. americana, C. heterophylla, C. colurna, and C. fargesii, were assessed for their response to field exposure to the eastern filbert blight (EFB) pathogen, Anisogramma anomala, in New Jersey, where the fungus is native. Plants were obtained from the U.S. Department of Agriculture Agricultural Research Service National Clonal Germplasm Repository and Oregon State University, the University of Nebraska, Lincoln, and the National Arbor Day Foundation. Additional plant material was acquired from the Morris and Holden Arboreta and from private nurseries in Amherst, NY, and Niagara-on-the-Lake, Ontario, Canada. The accessions were chosen based on their resistance to EFB in Oregon, a region where A. anomala is not native, or anecdotal reports and grower observations of tolerance or resistance to the disease. Trees were planted in the field from 2002 through 2009 in New Jersey where they were exposed to EFB yearly through field inoculations and natural spread. In Jan. 2012, they were visually evaluated for the presence of EFB. The cankers were measured, and the proportion of diseased wood was calculated for susceptible trees. Nearly all accessions reported to be resistant to EFB in Oregon maintained at least a useful level of tolerance in New Jersey with a number remaining free of cankers. However, several accessions developed small to medium-sized cankers and showed branch dieback, including offspring of C. avellana ‘Gasaway’. Most C. americana and C. heterophylla accessions remained free of EFB, although variation in EFB response was found in hybrids of these species with C. avellana, ranging from no signs or symptoms to severe EFB. Nearly half of the C. colurna × C. avellana hybrids developed cankers, whereas each of the C. fargesii accessions and most grower selections developed in eastern North America remained free of EFB. The results document the existence of a wide diversity of Corylus germplasm that expresses resistance or a high level of tolerance to EFB in New Jersey and confirms previous reports that C. americana is highly resistant to the disease. Interestingly, most C. heterophylla and the C. fargesii were also found to be resistant despite originating in Asia where A. anomala has not been found. The various interspecific hybrids show the potential for incorporating EFB resistance from wild species through breeding. The results provide further evidence of differences in disease expression in Oregon and New Jersey, where isolates differ and disease pressure may be higher.
John M. Capik and Thomas J. Molnar
Hazelnuts (Corylus sp.) are monoecious and wind-pollinated with reproduction limited by a sporophytic self-incompatibility system. They flower during the winter and are dichogamous with the dates of flowering ranging from December to March in New Jersey depending on the genotype, geographic location, and year. Successful, consistent nut production depends on both genetic compatibility and the appropriate timing of flowering between pollinizing and nut-producing cultivars. While the disease eastern filbert blight (EFB), caused by Anisogramma anomala, has severely limited past hazelnut production in the eastern United States, resistant and tolerant genotypes are now available for testing. However, little is known of their flowering phenology in this region. In this study, the flower and budbreak phenology of 19 different EFB-resistant and EFB-tolerant hazelnut accessions was evaluated over 4 years, and the results compared with air temperature data collected during bloom. Results showed that the accessions followed a similar progression of bloom each year (both staminate and pistillate flowers), which allowed their placement into early-, mid-, and late-flowering groups. However, the date of bloom and duration of bloom, especially for pollen shed, differed each year, largely corresponding to average air temperature trends. Confirming previous reports from other cold regions, it was shown that consistently colder average temperatures delayed bloom until later in the winter, which then led to a compressed period of flowering once temperatures warmed. In contrast, relatively warm temperatures over the season led to earlier flowering as well as a significant lengthening of the duration of bloom, similar to responses reported in Mediterranean climates. Our study documents hazelnut flowering phenology under New Jersey’s variable winter climate, and the results provide a benchmark for selecting suitable pollenizers and breeding parents for future nut production, flowering research, and/or genetic improvement in this region.
John M. Capik, Megan Muehlbauer, Ari Novy, Josh A. Honig, and Thomas J. Molnar
Stable genetic resistance to the fungal disease eastern filbert blight (EFB), caused by Anisogramma anomala, is vital for sustainable production of European hazelnut (Corylus avellana) in eastern North America. In this study, new hazelnut germplasm from the Russian Federation, Ukraine, and Poland (a total of 1844 trees from 66 seed lots) was subjected to A. anomala under field conditions over at least five years in New Jersey. Plants were then rated for the presence of EFB using an index of 0 (no disease) through 5 (all stems containing cankers). Nuts of the resistant trees were evaluated to identify plants with improved kernel characteristics. Genomic DNA of these trees was also screened with sequence-characterized amplified region (SCAR) markers generated by the primers BE-03, BE-33, and BE-68, which are closely linked to the single dominant R-gene of ‘Gasaway’, to assess the resistant seedlings for the presence of this well-known source of resistance. At final evaluation, 76 trees remained free of disease with nine expressing only minor symptoms (rating 1 or 2). The resistant trees spanned 24 different seed lots representing all three countries. The remaining trees ranged from moderately to severely infected with 81% of the total collection rating 5. Several of the resistant trees were found to produce commercial-sized (≈12 mm diameter), round kernels that blanched well. Although the results of the ‘Gasaway’ SCAR primers were inconclusive, the diverse collection origins and disease phenotypes provide evidence that novel sources of resistance were likely identified in this study. These new plants should broaden the genetic base of EFB-resistant C. avellana hazelnut germplasm available for breeding.
Thomas J. Molnar, Megan Muehlbauer, Phillip A. Wadl, and John M. Capik
Ronald S. Revord, Sarah T. Lovell, John M. Capik, Shawn A. Mehlenbacher, and Thomas J. Molnar
Eastern filbert blight (EFB), caused by the fungus Anisogramma anomala, is a primary limitation to european hazelnut (Corylus avellana) cultivation in eastern North America. American hazelnut (Corylus americana) is the endemic host of A. anomala and, despite its tiny, thick-shelled nuts, is a potentially valuable source of EFB resistance and climatic adaptation. Interspecific hybrids (Corylus americana × C. avellana) have been explored for nearly a century as a means to combine EFB resistance with wider adaptability and larger nuts. Although significant progress was made in the past, the genetic diversity of the starting material was limited and additional improvements are needed for expansion of hazelnut (Corylus sp.) production outside of Oregon, where 99% of the U.S. crop is currently produced. Our objective was to determine if C. americana can be a donor of EFB resistance. We crossed 29 diverse EFB-resistant C. americana accessions to EFB-susceptible C. avellana selections (31 total progenies) to produce 2031 F1 plants. In addition, new C. americana germplasm was procured from across the native range of the species. The new collection of 1335 plants from 122 seed lots represents 72 counties and 22 states. The interspecific hybrid progenies and a subset of the American collection (616 trees from 62 seed lots) were field planted and evaluated for EFB response following field inoculations and natural disease spread over seven growing seasons. EFB was rated on a scale of 0 (no EFB) to 5 (all stems containing cankers). Results showed that progeny means of the interspecific hybrids ranged from 0.96 to 4.72. Fourteen of the 31 progenies were composed of at least one-third EFB-free or highly tolerant offspring (i.e., ratings 0–2), transmitting a significant level of resistance/tolerance. Several corresponding C. americana accessions that imparted a greater degree of resistance to their hybrid offspring were also identified. In addition, results showed that 587 (95.3%) of the 616 C. americana plants evaluated remained completely free of EFB. These findings confirm reports that the species rarely expresses signs or symptoms of the disease and should be robustly studied and exploited in breeding.
Megan F. Muehlbauer, Josh A. Honig, John M. Capik, Jennifer N. Vaiciunas, and Thomas J. Molnar
The development of new cultivars resistant to the disease eastern filbert blight (EFB), caused by Anisogramma anomala, is of primary importance to hazelnut (Corylus sp.) breeders in North America. Recently, a large number of EFB-resistant cultivars, grower selections, and seedlings from foreign germplasm collections were identified. However, for a significant number of these, little is known of their origin, relationships, or genetic background. In this study, 17 microsatellite markers were used to investigate the genetic diversity and population structure of 323 unique accessions, including EFB-resistant and tolerant germplasm of uncertain origins, in comparison with a panel of known reference accessions representing a wide diversity of Corylus cultivars, breeding selections, and interspecific hybrids. The resulting allelic data were used to construct an unweighted pair group method using arithmetic averages (UPGMA) dendrogram and STRUCTURE diagram to elucidate relationships among the accessions. Results showed 11 consensus groups with EFB-resistant or tolerant accessions in all, providing strong evidence that EFB resistance is relatively widespread across the genus Corylus. Furthermore, open-pollinated seedlings tended to group together with reference accessions of similar geographic origins, providing insight into their genetic backgrounds. The results of this study add to the growing body of knowledge of hazelnut genetic resources and highlight recently introduced EFB-resistant seedling germplasm as new, unrelated genetic pools of resistance.
Josh A. Honig, Megan F. Muehlbauer, John M. Capik, Christine Kubik, Jennifer N. Vaiciunas, Shawn A. Mehlenbacher, and Thomas J. Molnar
European hazelnut (Corylus avellana L.) is an economically important edible nut producing species, which ranked sixth in world tree nut production in 2016. European hazelnut production in the United States is primarily limited to the Willamette Valley of Oregon, and currently nonexistent in the eastern United States because of the presence of a devastating endemic disease, eastern filbert blight (EFB) caused by Anisogramma anomala (Peck) E. Muller. The primary commercial means of control of EFB to date is through the development and planting of genetically resistant european hazelnut cultivars, with an R-gene introduced from the obsolete, late-shedding pollinizer ‘Gasaway’. Although the ‘Gasaway’ resistance source provides protection against EFB in the Pacific northwestern United States (PNW), recent reports have shown that it is not effective in parts of the eastern United States. This may be in part because the identification and selection of ‘Gasaway’ and ‘Gasaway’-derived cultivars occurred in an environment (PNW) with limited genetic diversity of A. anomala. The objectives of the current research were to develop a genetic linkage map using double digestion restriction site associated DNA sequencing (ddRADseq) and identify quantitative trait loci (QTL) markers associated with EFB resistance from the resistant selection Rutgers H3R07P25 from southern Russia. A mapping population composed of 119 seedling trees was evaluated in a geographic location (New Jersey) where the EFB fungus is endemic, exhibits high disease pressure, and has a high level of genetic diversity. The completed genetic linkage map included a total of 2217 markers and spanned a total genetic distance of 1383.4 cM, with an average marker spacing of 0.65 cM. A single QTL region associated with EFB resistance from H3R07P25 was located on european hazelnut linkage group (LG) 2 and was responsible for 72.8% of the phenotypic variation observed in the study. Based on its LG placement, origin, and disease response in the field, this resistance source is different from the ‘Gasaway’ source located on LG6. The current results, in combination with results from previous research, indicate that the H3R07P25 source is likely exhibiting resistance to a broader range of naturally occurring A. anomala isolates. As such, H3R07P25 will be important for the development of new european hazelnut germplasm that combines EFB resistance from multiple sources in a gene pyramiding approach. Identification of EFB resistance in high disease pressure environments representing a diversity of A. anomala populations is likely a requirement for identifying plants expressing durable EFB resistance, which is a precursor to the development of a commercially viable european hazelnut industry in the eastern United States.
David E. Zaurov, Thomas J. Molnar, Sasha W. Eisenman, Timothy M. Ford, Ravza F. Mavlyanova, John M. Capik, C. Reed Funk, and Joseph C. Goffreda
Central Asia is a center of diversity for many important fruit and nut tree species, including wild and cultivated apricots (Prunus armeniaca L.). A wealth of apricot germplasm that expresses novel and valuable characteristics such as fruits with high soluble solids, edible kernels, glabrous skin, and diverse colors and flavors, as well as later-blooming flowers, late-maturing fruit, and drought, cold, and salt tolerance, can be found growing across this region. Since the dissolution of the Soviet Union, Central Asia has become more accessible for reciprocal germplasm exchange and scientific collaborations. Thus, opportunities now exist to obtain, study, and use a much wider diversity of Central Asian apricot germplasm in breeding efforts, which can lead to improved crop traits and ultimately an expansion of the regions where this high-value crop can be grown. To bring attention to the valuable P. armeniaca genetic resources found in Central Asia and to promote its better use, management, and preservation, a description and history of the species from a Central Asian perspective, along with recent and ongoing activities, are discussed in this article.