Hymenoscyphus fraxineus, an introduced ascomycete fungus and primary causal agent of European ash dieback, was investigated on Fraxinus mandshurica trees in its native range in Primorye region of Far East Russia. This evidence is the first report of H. fraxineus on healthy, asymptomatic F. mandshurica trees. High-throughput sequencing revealed 49 distinct fungal taxa associated with leaves of F. mandshurica, 12 of which were identified to species level. Phyllosphere fungal assemblages were similar among sites despite being largely geographically distant. Many organisms comprising the foliar fungal community on F. mandshurica in Far East Russia have similarity to those reported inhabiting F. excelsior in Europe based on previous studies. However, Mycosphaerella sp., the most dominant species in this study and detected in nearly all samples, was associated only with F. mandshurica. Genetic diversity of H. fraxineus was significantly higher in the Far East Russian population than in Europe. In contrast to its aggressive behaviour on Fraxinus excelsior in Europe, H. fraxineus appears to be a benign associate of indigenous F. mandshurica that initially induces quiescent and asymptomatic infections in healthy trees prior to active host colonization normally associated with modification of host tissue during senescence.
Ten juvenile Fraxinus excelsior half-sib families from two Lithuanian populations have been tested in the controlled environment for their response to ash dieback disease caused by Hymenoscyphus fraxineus, detecting changes of genetic variation and heritability, as well as estimating genotype by environment (G×E) interaction and phenotypic plasticity following artificial spring frost and summer drought treatments. In 2014, a batch of 200 four-year-old ash seedlings was used for each treatment and control (no treatment). Health condition, bud flushing phenology and height were assessed for each seedling, and disease incidence and survival ratios were assessed for each family both before (at the beginning of the vegetation season) and after the treatments (at the end of the vegetation season). Disease incidence ratio increased from 0.77-0.80 up to 0.90-0.95. Tree mortality rates during one vegetation season were significantly lower in the frost treatment (21%) than in the drought treatment (25%) or control (31%). None of the tested F. excelsior families were completely resistant to ash dieback, although significant among-family differences in disease incidence and damage rates suggest an additive mode of gene action and thus a quantitative resistance to the disease. Neither disease incidence rates, nor tree health condition scores differed significantly among the applied treatments (including control) indicating in general a negligible effect of the simulated adverse conditions on health status of the ash seedlings. However, G×E interaction was found to be significant (at P > 0.05) for disease incidence, length of necrotic shoots and tree survival, implying that susceptibility of ash families to the dieback disease unequally depends on environmental conditions, and indicating a presence of genetic variation in plasticity and reaction norms of the tested families across environments (treatments). Substantially increased coefficients of additive genetic variation and heritability in health condition following both frost and drought treatments and compared to control showed that simulated stress conditions may noticeably contribute to expression of differences among the tested F. excelsior families in their resistance traits, thus enabling a better evaluation of performance of different families, an effective family selection for resistance, and achievement of a marked genetic gain.
Following its introduction from Asia in the 1990s, the ascomycete Hymenoscyphus fraxineus has caused a severe dieback of Fraxinus excelsior in Europe. In this study, the virulence of 200 H. fraxineus isolates were assessed and compared. These isolates equally represented (i) two geographically distant populations with a different disease history (Switzerland, recently established populations at the epidemic front versus Lithuania, old established populations), and (ii) isolates from two different types of host tissue (necrotic bark lesions as dead‐end tissue versus fallen leaf petioles as primary host tissue). Inoculations conducted on 3‐year‐old F. excelsior seedlings showed that the vast majority of the isolates (98%) were able to induce necrotic bark lesions after 10 months. Although a high variation in virulence was observed among isolates, no significant differences were detected between the older and the epidemic‐front populations. Decline in virulence of populations of invasive organisms is generally assumed with increasing age of epidemics. However, this does not appear to hold true for H. fraxineus. Either the Lithuanian population is still too young (15 to 20 years old) to show a decline in virulence, or the size of the host population may still not be critical for pathogen survival. Given that bark lesions represent an epidemiological dead end and do not benefit the survival of H. fraxineus, a trend towards reduced ‘bark virulence’ of isolates originating from leaf petioles compared to isolates from the bark lesions was expected. However, such a trend was observed neither in old, nor in recently established populations.
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