The moisture content of timber in relation to attack by Lyctus powder-post beetles

The relation between moisture content of oak sapwood and rate of development of Lyctus brunneus has been investigated at 23C. Oviposition took place normally in samples of various moisture contents between 5.5 and 23.5%. The eggs hatched in wood of 7.7% moisture content, but not of 61% moisture content, although the embryos developed fully. The rate of gain in weight of young larvae increased to a maximum as the moisture content of the wood rose to about 14.2% and then fell again. The rate of development from egg to adult followed a somewhat similar curve with a maximum at 16% moisture content. At all moisture contents the first male emerged a few days earlier than the first female. The average weight of the male beetles was less and their moisture content greater than those of the corresponding females. The average weight of the beetles showed a tendency to increase at moisture contents of the wood above 97%. The average moisture content of the males was fairly constant up to 156% moisture content of the wood, but that of the females rose slowly as the moisture content of the samples increased. In wood of 15.6% moisture content at 23C. the minimum period of development from egg to adult was 167 days; at 14.2% moisture content and 27.5 C. this was reduced to 142 days. Green timber was shown to be liable to infestation as soon as a superficial skin was formed of a suitable moisture content; the upper limiting moisture content is probably the fibre saturation point. Control of moisture content of timber cannot be achieved sufficiently easily for it to become an important method for the control of Lyctus infestation.     

Micropropagation of common ash (Fraxinus excelsior)

Embryos extracted from dried seeds of common ash (Fraxinus excelsior), were germinated on growth regulator-free culture medium. Cotyledonary nodes from these seedlings were placed onto Murashige and Skoog, Woody Plant or Driver and Kuniyuki culture media with 22.2 or 44.4 M benzyladenine, on which they developed into shoot cultures following the outgrowth of axillary buds. With Murashige and Skoog medium, cultures often died. With Woody Plant Medium, survival of the cultures was considerably improved, but large amounts of callus were produced at the cut ends of the explants, and new axillary shoots had long internodes and small leaves. With Driver and Kuniyuki medium, both survival and callus formation were much improved, and the shoots produced were of high quality. Proliferation of axillary shoots was obtained from both shoot tip and nodal explants placed onto Driver and Kuniyuki medium with 22.2 M benzyladenine. Adventitious root formation was best with shoots inserted into half-strength Woody Plant Medium containing 2.45, 4.9 or 9.8 M indolebutyric acid. All of the rooted plantlets tested have successfully established in soil.     

Micropropagation of hornbeam (Carpinus betulus L.) and ash (Fraxinus excelsior L.)

Shoot multiplication of hornbeam was stimulated onWPM, QL andDKW medium supplemented with a low concentration of BAP or BPA (0.1-0.2 mg I -1) andIBA (0.1 mg I -1). Low concentration of thidiazuron promoted axillary bud formation, higher concentration inhibited shoot elongation. Microshoots were rooted onWPM supplemented with a low auxin concentration (IBA or NAA 0.2-0.5 mg I -1). High rooting percentages were obtained. Shoot proliferation of ash was stimulated on MS andDKW medium supplemented withBAP orBPA (2.0-5.0 mg I -1) andIBA (0.1 mg I -1). Root formation was promoted onWPM containing a low auxin concentration. Rooted plantlets were transplanted into soil and after hardening off the micropropagated trees were planted in the field. The planted trees grew normally without showing signs of abnormality.     

Regeneration of shoots from embryo hypocotyls of common ash (Fraxinus excelsior)

Summary The addition of thidiazuron (TDZ) to MS salts and vitamins, instead of benzylaminopurine (BAP) increased both the culture weight and the proportion of ash embryo hypocotyl explants that produced adventitious shoots. The concentration of BAP, but not that of TDZ also affected these parameters. Addition of 1-naphthalene acetic acid reduced culture performance, indolebutyric acid (IBA) was beneficial, and 2,4-dichlorophenoxyacetic acid had no effect. During both 1990 and 1991, rates of regeneration declined as the growth season progressed. Adventitious shoots, which were also obtained from hypocotyls from dried seeds, were established as proliferating shoot cultures following transfer to DKW medium with 5.0 mg l^–1 BAP, and the resulting shoots were rooted in half-strength Woody Plant medium with 1.0 mg l^–1 IBA.     

Frost periods and beginning of the ash (Fraxinus excelsior L.) pollen season in Basel (Switzerland)

In order to be able to forecast the beginning of the ash pollen season, several meteorological factors have been considered. During these research activities it appeared that the ash pollen release is closely linked to frost periods happening during the months of February and March. For 24 years over a period of 29 consecutive years, a daily concentration of 30 ash pollen grains/m³/day has been reached when the cumulated mean daily temperatures starting on the last day when the absolute minimum temperature is lower or equal to −2 °C amounts on average to 214 °C – admitting a variation of plus or minus 42 °C. As for the five remaining years, rain is three times responsible for the delay of the pollen release. This revised version was published online in June 2006 with corrections to the Cover Date.     

Factors affecting the survival of ash (Fraxinus spp.) trees infested by emerald ash borer (Agrilus planipennis)

Emerald ash borer (Agrilus planipennis) (EAB), an Asian woodboring beetle accidentally introduced in North America, has killed millions of ash (Fraxinus spp.) trees and is spreading rapidly. This study examined the effects of tree- and site-level factors on the mortality of ash trees in stands infested by EAB in OH, USA. Our data show that ash populations in forested sites can progress from healthy to almost complete mortality of mature trees within 6 years. Although the end result of nearly complete mortality does not vary, survival analysis with 5 years of data showed that some factors affected the rate of mortality. We found more rapid mortality in stands with lower densities of ash trees. This finding supports an extension of the resource dilution hypothesis whereby concentration of EAB on few trees in low ash density areas leads to rapid decline of these trees. This contradicts an extension of the resource concentration theory that greater host density increases relative pest abundance and host mortality. Although reductions in ash density via diversification may be desirable for other silvicultural, conservation, and management objectives in preparation for EAB, our study shows that the management strategy of reducing ash density is unlikely to protect the remaining ash trees. Survival analysis also showed that mortality was more rapid for trees shaded by other trees and for trees initially exhibiting dieback. In management scenarios where hazard tree removal must be spread over several years due to budget constraints, focusing initial tree removal on stressed trees is recommended.     

Underwater germination potential of common ash seed (Fraxinus excelsior L.) originating from flooded and non-flooded sites

Oxygen availability is one of the main factors affecting seed germination capacity. On floodplains, oxygen supply is seasonally restricted because of flooding events. Seed ecotypes are able to cope with total submergence to different extents, depending on the species and its position in a given environmental gradient. This investigation tested three provenances of common ash (Fraxinus excelsior L.), one from a lowland site and two from upland sites, for germination capacity and germination velocity under water. The final germination capacity of all provenances was unaffected, but the germination velocity was decreased for all provenances. The adaptational significance of these results is discussed.     

Cumulative temperatures for prediction of the beginning of ash (Fraxinus excelsior L.) pollen season

The aim of this study is the analysis of the meteorological conditions leading to the release of ash pollen in order to predict the beginning of the pollen season in Zurich, Switzerland. For 12 years over a period of 14 consecutive years, it appears that ash pollen release begins when the cumulated mean daily temperatures starting on January 1 reach 220.7°C, admitting a variation of ±30°C that more or less corresponds to 3 consecutive days at 10°C.     

Chloroplast DNA variation and postglacial recolonization of common ash (Fraxinus excelsior L.) in Europe

We used chloroplast polymerase chain reaction-restriction-fragment length polymorphism (PCR-RFLP) and chloroplast microsatellites to assess the structure of genetic variation and postglacial history across the entire natural range of the common ash (Fraxinus excelsior L.), a broad-leaved wind-pollinated and wind-dispersed European forest tree. A low level of polymorphism was observed, with only 12 haplotypes at four polymorphic microsatellites in 201 populations, and two PCR-RFLP haplotypes in a subset of 62 populations. The clear geographical pattern displayed by the five most common haplotypes was in agreement with glacial refugia for ash being located in Iberia, Italy, the eastern Alps and the Balkan Peninsula, as had been suggested from fossil pollen data. A low chloroplast DNA mutation rate, a low effective population size in glacial refugia related to ash's life history traits, as well as features of postglacial expansion were put forward to explain the low level of polymorphism. Differentiation among populations was high (GST= 0.89), reflecting poor mixing among recolonizing lineages. Therefore, the responsible factor for the highly homogeneous genetic pattern previously identified at nuclear microsatellites throughout western and central Europe (Heuertz et al. 2004) must have been efficient postglacial pollen flow. Further comparison of variation patterns at both marker systems revealed that nuclear microsatellites identified complex differentiation patterns in south-eastern Europe which remained undetected with chloroplast microsatellites. The results suggest that data from different markers should be combined in order to capture the most important genetic patterns in a species.     

Genetic variability of Chalara fraxinea, dieback cause of European ash (Fraxinus excelsior L.)

A total of 159 colonies of Chalara fraxinea were isolated between 2005 and 2006 from dying trees of European ash (Fraxinus excelsior L.) aged between 3 and 10 years. They derived from five regions of Poland differing by geographic location and climatic conditions. On the basis of 90 RAMS markers, pathogen intra- and inter-population variability, as well as its dependency on geographic distance and climatic conditions in the regions of strain origin, was analysed. The applied measures of intrapopulation genetic variability (genetic distance, Nei’s unbiased diversity, Shannon’s Information Index and percentage of polymorphic loci) allowed for differentiation of two strain groups: the first deriving from lowlands and the second from uplands and mountainous areas. Strains in lowlands were characterised by smaller number of markers, smaller number of polymorphic loci and smaller intrapopulation genetic variability. Positive and statistically significant correlation was shown between variability of isolates and elevation of regions above sea level. Pair-wise genetic distances between groups of isolates (Nei’s unbiased genetic distance) from particular regions were not significantly correlated with the corresponding geographic distances. On the basis of AMOVA, it was shown that 85% of variability was within-region differences and 2% between-region differences, whereas differences between lowlands and uplands were 13%. Principal Components Analysis (PCA) for the investigated regions confirmed the results from Nei’s genetic distance matrix.     

Assessment of genetic structure within and among Bulgarian populations of the common ash (Fraxinus excelsior L.)

We analysed genetic variation within and between populations of the common ash from Bulgaria in order to extract biological information useful in the context of conservation management of eastern European genetic resources of noble hardwood species. A total of 321 trees from three regions of Bulgaria were typed at six highly polymorphic microsatellite loci. Analysis of within-population inbreeding suggests an upper boundary value of 2.7% for the selfing rate. Significant spatial genetic structure consistent with models of isolation by distance was detected within four out of 10 populations as well as among populations. Estimates of neighbourhood size in the range 38-126 individual trees were obtained based on spatial genetic structure analyses at either the intrapopulation or interpopulation level. Differentiation among populations explained only about 8.7% of total genetic diversity. These results are discussed in comparison with data from social broad-leaved species such as oak and beech.     

Nitrogen storage and remobilization in ash (Fraxinus excelsior) under field and laboratory conditions

 Storage and remobilization of nitrogen (N) were studied in ash trees (Fraxinus excelsior) under both field and greenhouse conditions. Experiments in the greenhouse providing ¹⁵N labelled fertilizer to the trees showed that the major quantity of N remobilized during subsequent spring was from the roots, and only a small amount from the stem. This corresponded with a loss of soluble N (proteins and low-molecular-weight compounds) from both roots and stem. On the two field sites, which differed in water availability, there was a decrease in bark N content during leaf growth, but on the dry site net N export from the bark was sustained throughout the whole vegetation period. Remobilized N was derived from soluble proteins and low-molecular-weight compounds on the moist site, which was demonstrated by the seasonal dynamics of a 56 kDa polypeptide in bark and wood. On the dry site, lower contents of soluble proteins were associated with smaller amounts of N remobilized compared to the moist site. Uptake studies of ¹⁵N labelled fertilizer indicated a higher contribution of current uptake to leaf N increment during spring at the dry site compared to the moist site. Differential N availability during the season had a decisive effect on the nitrogen storage dynamics at the two sites. Thus the influence of current N supply on N remobilization and storage as found in the greenhouse-grown plants could be verified under field conditions. Received: 28 July 1995 / Accepted: 17 July 1996     

Spatio-temporal dynamics of endophyte diversity in the canopy of European ash (Fraxinus excelsior)

Leaf-inhabiting endophytic fungi of Fraxinus excelsior growing in a floodplain forest were isolated during 2008 to investigate vertical community structure, species richness and seasonal variation. The analysis of 848 fungal endophytes from 213 leaves resulted in 50 different species. In the understorey, infection density and species richness were higher than in the crowns of mature trees throughout the whole vegetation period. Within tree crowns, sun-exposed leaves of the top canopy exhibited the lowest infection rates. Most species were rare or absent in spring and in the light crowns and frequent in autumn and the understorey. However, some species, especially the two most frequent, Alternaria infectoria and A. alternata, deviated from these patterns. Young leaves were nearly free of endophytes. Apparently, the subsequent infection and establishment of fungi strongly depend on microclimatic parameters and leaf characters, which create highly variable spatial and temporal colonisation patterns within an individual tree.     

Effects of grafting on host plant resistance in ash (Fraxinus spp.) to emerald ash borer (Agrilus planipennis Fairmaire)

1. The invasive emerald ash borer (EAB) (Agrilus planipennis Fairmaire Coleoptera: Buprestidae) is a phloem‐boring pest from Asia that has killed hundreds of millions of North American ash trees (Fraxinus spp.). Propagation of trees resistant to EAB must be explored as a tool to maintain the North American ash resource.
2. Interactions between the root and shoot of a tree may be critical to its ability to resist attack by insects and pathogens because many secondary metabolites critical to plant defence are synthesized in the roots.
3. In the present study, we determined the extent to which grafting susceptible green ash (Fraxinus pennsylvanica Marshall) and resistant Manchurian ash (Fraxinus mandshurica Rupr.) affects EAB resistance in ash trees. We measured adult EAB survivorship, herbivory, fecundity and morphological tree characteristics on grafted and nongrafted green and Manchurian ash.
4. Among grafted trees, beetles caged on grafted green ash lived four days longer and laid at least three times more eggs than beetles caged on trees with a Manchurian ash scion or rootstock. Herbivory was unaffected by graft combination.
5. Our findings suggest that grafted trees with Manchurian ash rootstocks or scions will be more resistant to EAB than nongrafted green ash trees.

     

Interspecific variation in resistance to emerald ash borer (Coleoptera: Buprestidae) among North American and Asian ash (Fraxinus spp.)

We conducted a 3-yr study to compare the susceptibility of selected North American ash and an Asian ash species to emerald ash borer, Agrilus planipennis Fairmaire, an invasive wood-boring beetle introduced to North America from Asia. Because of a coevolutionary relationship between Asian ashes and emerald ash borer, we hypothesized an Asian ash species, Manchurian ash, is more resistant to the beetle than its North American congeners. Consistent with our hypothesis, Manchurian ash experienced far less mortality and yielded far fewer adult beetles than several cultivars of North American green and white ash. Surprisingly, a black ash (North American) x Manchurian ash hybrid was highly susceptible to emerald ash borer, indicating this cultivar did not inherit emerald ash borer resistance from its Asian parent. A corollary study investigated the efficacy of soil-applied imidacloprid, a systemic, neonicotinoid insecticide, for controlling emerald ash borer in each of the five cultivars. Imidacloprid had no effect on emerald ash borer colonization of Manchurian ash, which was low in untreated and treated trees. In contrast, imidacloprid did enhance survival of the North American and hybrid cultivars and significantly reduced the number of emerald ash borer adults emerging from green and white ash cultivars. We identify a possible mechanism of resistance of Manchurian ash to emerald ash borer, which may prove useful for screening, selecting, and breeding emerald ash borer-resistant ash trees.