A review of exapted resistance to diseases of Populus

The cultivation of hybrid poplar in the Pacific North-west (PNW) has expanded in the last 15 years. Native Populus trichocarpa is susceptible to many pests and fungal pathogens but its interspecific hybrids were largely disease-free until 1991 when Melampsora medusae f. sp. deltoidae first appeared in the region. Genetic analyses of resistance in the PNW, and elsewhere, reveal a pattern whereby genes for resistance are inherited in interspecific hybrids from the exotic, noncoevolved Populus parent. With some pathogen/poplar hybrid combinations, this resistance segregates in the F₁ generation, but with others, only in the F₂. Such exapted resistance is now known to occur in P. trichocarpa (versus the exotic, eastern North American rust species, M. medusae f. sp. deltoidae), in Populus deltoides (versus the Eurasian rust, Melampsora larici-populina, the PNW population of Septoria populicola, and an undescribed PNW eriophyid mite), and in Populus maximowiczii (versus Melampsora occidentalis, and the PNW populations of Venturia populina, S. populicola, and Linospora tetraspora).     

Development and use of new sensitive molecular tools for diagnosis and detection of Melampsora rusts on cultivated poplar

Poplar rusts due to Melampsora larici‐populina (Mlp), M. allii‐populina (Map) and M. medusae f. sp. deltoidae (Mmd) are the most serious disease in Europe on cultivated poplars, that is, Populus × euramericana and P. × interamericana hybrids. These pathogenic species can be identified by the observation of morphological characteristics of urediniospores but this method is not appropriate for high‐throughput analysis and cannot be used on other spore stages, such as aeciospores or teliospores, that are morphologically similar. The aim of this study was to develop a rapid and sensitive molecular method based on PCR amplification that was able to specifically detect these species on various hosts for routine analysis. Three primer pairs ITS‐MLP‐F/ITS‐MLP‐R, ITS‐MAP‐F/ITS‐MAP‐R and ITS‐MMD‐F/ITS‐MMD‐R were designed within the internal transcribed spacer (ITS) sequences of ribosomal DNA to target Mlp, Map and Mmd, respectively, and their specificity were confirmed on a wide range of isolates and species. ITS‐MLP‐F/ITS‐MLP‐R and ITS‐MAP‐F/ITS‐MAP‐R primers proved to be highly specific to Mlp and Map, respectively, whereas ITS‐MMD‐F/ITS‐MMD‐R cross‐reacted with DNA from M. larici‐tremulae and M. pinitorqua. However, these species are not pathogenic on cultivated poplars that all belong to sections Aigeiros and Tacamahaca of the genus Populus. Specific Mmd primers proved to be very sensitive as a positive signal could be obtained with DNA extracts from 6 target urediniospores mixed with 800 000 urediniospores of Mlp. An internal amplification control (IAC) was included to discriminate false negative results due to the potential presence of inhibitory compounds in DNA extracts. ITS‐MMD‐F/ITS‐MMD‐R primers are therefore efficient for the detection of the quarantine pathogen Mmd on samples collected on poplar or larch and are fit for use in official tests. This new PCR assay has been used in routine for ten years, and Mmd has hitherto never been detected in commercial poplar nurseries in France.     

The detection of poplar mosaic virus and its occurrence in a range of clones in England and Poland

The incidence of poplar mosaic virus evidenced by foliage symptoms, enzyme-linked immunosorbent assay (ELISA) or bioassay (infectivity) was greater in stooled clones having Aigeiros (especially as the female) parents than in those of Tacamahaca parentage. The overall prevalence of the virus in clones growing in England was less (84/344, 24%) than in clones growing in Poland (143/256, 56%). Whereas symptoms in PMV-infected P. deltoides were prominent and common, in P. nigra, another species assigned to the Section Aigeiros they were indistinct and rare. Evidence of poplar mosaic virus inferred from ELISA alone was found in only one aspen of 52 tested clones in the Sections Leuce, Leuce × Leuce or Leucoides.     

Climatic favourability zones for Eucalyptus rust in Brazil

Brazilian forest‐based industries are supported by more than 5.5 million hectares of Eucalyptus growing under different climatic conditions with different degrees of favourability for rust Puccinia psidii, including both traditional and expanding areas, where such disease is of major concern for the industry, foresters and scientists. The main objective of this study was to define favourable climatic zones for Eucalyptus rust in Brazil with the following aims: (i) to develop a spatial method for estimating the mean night‐time temperature (T_ng); (ii) to assess and validate a Eucalyptus rust model; and (iii) to map Eucalyptus rust favourability zones in Brazil based on the proposed model. A straightforward method, based only on latitude, day of the year, maximum and minimum air temperatures, was developed to estimate T_ng, which is a key variable for a proper application of Ruiz rust model. Based on 37 field experiments with natural rust occurrence, it was observed that climatic conditions are determining factors for disease severity. Significant correlations between disease severity observed in the field and climatic conditions were found: a negative correlation with temperature (r = .50, p < .01) and a positive correlation with relative humidity (r = .89, p < .0001). A significant correlation (r = .81, p < .0001) between the normalized infection index, produced from Ruiz model, and the average rust score was also observed. Once the model was validated under field conditions, it was applied using historical average data of air temperature and leaf wetness duration to obtain monthly Eucalyptus rust favourability maps for the whole country. These final maps show that favourable climatic zones for Eucalyptus rust are extremely dynamic, with high temporal and spatial variability in Brazil and that climatic conditions should be considered for expansion of Eucalyptus to new areas, in breeding programmes, and for defining the most suitable seasons for forest establishment in each climate of the country. These results provide forestry managers with practical tools to reduce uncertainty about the expected severity of Eucalyptus rust in Brazil.     

Improvement of wood properties by urea-formaldehyde resin and nano-SiO2

In order to improve wood properties of triploid clones of Populus tomentosa, urea-formaldehyde (UF) resin was compounded with nano-SiO₂, coupling agents and flame retardants in different ways to prepare five kinds of modifiers. The poplar wood samples were impregnated with the modifiers and heated to prepare UF-SiO₂-wood composites. The antiswelling efficiency, resistance of water absorption, oxygen index and hardness of the composites were measured. Results show that all of the modifiers reduced water absorption of poplar wood and enhanced flame resistance and hardness. Nano-SiO₂ showed a marked effect in improving the hardness of wood. In addition, all of the modifiers, except UF-C-SiO₂-polymer, improved the dimensional stability of poplar wood. The UF resin and nano-SiO₂ compound improved general properties of poplar wood. __________ Translated from Journal of Beijing Forestry University, 2006, 28(2): 123–128 [译自: 北京林业大学学报]     

Variation in Melampsoridium resistance among European white-birch clones grown in different fertilization treatments

Resistance of 2-year-old plantlets of seven European white birch, Betula pendula, clones to birch rust, Melampsoridium betulinum, was studied in field experiments and in a leaf-disc bioassay. In addition, rust resistance of plantlets growing in a nursery under three fertilization treatments was tested. The birch clones clearly varied in their levels of resistance. One of the clones was consistently the most resistant, and two were very susceptible. Plantlets growing in the lowest fertility treatment were the most resistant. Clone × fertilization interaction was small. The plantlets grown in the lowest fertilization treatment, in particular, deviated from the other treatments. Generally good rust-resistance correlations were obtained between different experiments. The leaf-disc bioassay was an effective way of determining the field rust resistance of birch clones. The possibility of trade-offs between rust resistance and tree growth is discussed.     

Effect of Melampsora medusae leaf rust infection on yield of hybrid poplars in the north-central United States

Melampsora medusae Thuem. is a hazard to cultivation and fiber production of hybrid poplars grown under short rotations, particularly when the alternate host is present. This rust can be controlled with biweekly fungicide applications but resistant clones are the best method of control.     

Infection of Populus alba var. hickeliana by Melampsora medusae Thüm.

The first record of infection of Populus alba var. hickeliana by the American rust, Melampsora medusae Thum. is reported. The morphology of uredospores of rust on P. alba is compared with that of other Melampsora spp. infecting white poplars. The implications of these observations for the widespread establishment in the future of white × black hybrid poplars are discussed.     

CONTROL TECHNIQUE OF BACTERIAL CANKER OF POPLAR

The poplar bacterial canker caused by Erwinia sp is a serious disease spreading inmost parts of Northeast China,where poplars are cultivalted.The disease seriously infects thosepoplars belonging to section Aigeiros or the intersection hybrids of section Tacamahaca and section Aigeiros and their clones.Five fine poplar cultivars resistant to the bacterial canker have been ob-tained by screening tests.They are Lz_2(Populus simonii×nigra var.italica),A_(15)(P.×sp.)613(P.×sp.),A_(102)and A_(98)(P.simonii×sp.)T following chemicals are obviously effective,such as:50％DT germicide ,40％ XF-136 and 10％ C.C.M.A.or 12.5％ ramphencol and 50,000 ppm/mlvalidamycin,with which the stems or roots can be watered.     

Alder rust,Melampsoridium hiratsukanum Ito,identified from Wales,UK and British Columbia,Canada

An epidemic rust, Melampsoridium hiratsukanum, originating from East Asia has spread to Europe over the last two decades. We report the first observation of this rust in the British Isles on leaves of Alnus incana and A. glutinosa. In addition to M. hiratsukanum, a native M. betulinum–like rust co‐occurred on the same leaves as a mixed infection. We also confirm that a rust on A. rubra in Western Canada is identical to M. hiratsukanum present in East Asia and Europe.     

Effect of environment on the response of Eucalyptus clones to inoculation with Cryphonectria cubensis

The ascomycete Cryphonectria cubensis causes severe losses in Eucalyptus plantations in South Africa and selection programmes for disease tolerance are necessary. The aim of this study was to use two C. cubensis inoculation trials, planted at different locations to assess the disease susceptibility of the clones and the effect of the environment on disease development. These two trials consisted each of 21 Eucalyptus clones (E. grandis, E. grandis × camaldulensis and E. grandis × urophylla). All trees were inoculated with a single virulent strain of C. cubensis and lesion widths measured 6 and 12 months after inoculation. Clones differed significantly in their tolerance to C. cubensis. Further, disease severity differed depending on the geographical location of the trial. A significant clone × locality (genotype × environment) interaction was observed. Therefore, screening for disease resistance should take place only in areas where clones will be commercially grown.     

An assessment of potential of hybrid poplar for planting in the Virginia Piedmont

Poplar species grow well across the temperate zone, but hybrid varieties have not previously been evaluated for planting in the Virginia Piedmont region. The top 12 clones in height growth and rust resistance from a screening trial involving 98 hybrid poplar varieties of three taxa (Populus deltoides × P. maximowiczii, DxM; P. deltoides × P. nigra, DxN; P. deltoides × P. trichocarpa, DxT) were selected for planting in replicated yield trials at two locations in the Virginia Piedmont. Results through the first four growing seasons showed that the DxM taxon had the most rapid height development. It was, however, the taxon most affected by a late spring frost at the Appomattox-Buckingham State Forest site and by Septoria stem canker at the Reynolds Homestead site. Analysis of variance of clonal and location effects showed highly significant differences among replicates within location and among clones within taxon. Among seven clones within the DxM taxon, pairwise comparison tests of height growth identified two groups: a group of four better clones that were significantly different (p = 0.05) from a second group of three. These early results suggest that multi-selection criteria, including growth, disease and frost resistance, are important when developing hybrid poplar clones for planting in the Piedmont region.     

Identification and characterization of fungi associated with internal wood lesions and decline disease of willow and poplar trees in Iran

During the study of fungal trunk pathogens associated with urban trees decline in Shiraz (Iran), a serious decline of willow and poplar trees was observed. Therefore, an investigation was conducted on these trees in some areas of this city during spring and summer 2012 and 2013, to determine the main fungal trunk pathogens associated with these ornamental plants. Plant materials were collected from trees exhibiting disease symptoms such as yellowing, shoot canker, shoot dieback, defoliation and internal wood necrosis and decayed wood. Fungal isolations were made from discoloured or decayed wood tissue onto 2% malt extract agar (MEA) amended with streptomycin sulphate. Nine species, Fomes fomentarius, Diplodia seriata, Lasiodiplodia theobromae, Dothiorella sarmentorum, Neoscytalidium hyalinum, Diatrype whitmanensis, Phaeoacremonium rubrigenum, P. aleophilum and P. parasiticum, were identified based on morphology and DNA sequence comparisons. Pathogenicity tests were performed on detached shoots of willow and poplar trees under greenhouse conditions. Lasiodiplodia theobromae caused the longest lesions on willow. On poplar shoots, the longest lesions were caused by P. parasiticum. Diplodia seriata produced the smallest lesions on both woody hosts. First reports from willow wood include P. parasiticum, P. rubrigenum, D. whitmanensis, L. theobromae, D. seriata and N. hyalinum, while new reports from poplar wood include P. parasiticum and Do. sarmentorum. Based on our knowledge, this is also the first report of D. whitmanensis in Iran.     

Melampsora and Marssonina pathogens of poplars and willows in New Zealand

New Zealand has hosted three Melampsora species attacking poplars (Populus spp.), namely, Melampsora larici-populina, Melampsora medusae and for a single season, Melampsora medusae-populina, a unique interspecific hybrid. The predominant species, M. larici-populina, over-winters via larch and each year new races appear causing defoliation of previously rust-resistant cultivars. Melampsora medusae also occurs in some seasons following over-wintering on certain hosts or by aerial transport from Australia. Willows (Salix spp.) host four Melampsora species, Melampsora coleosporioides, Melampsora epitea var. epitea and two unidentified species attacking Salix viminalis and Salix daphnoides/Salix incana× open pollinated hybrids, respectively. The host range and pathogenicity of these species have remained static as no breakdown in host resistance has been observed. The anthracnose fungi, Marssonina brunnea and Marssonina castagnei on poplar, and Marssonina salicicola on tree willows occur throughout New Zealand. All three species can cause severe defoliation of susceptible cultivars in moist districts where annual rainfall exceeds 1000 mm. Although all form abundant microconidia during autumn, the Drepanopeziza states have never been observed in New Zealand. During an unusually wet summer (199–93), a new strain of Marssonina salicicoa appeared which defoliated the previously resistant cultivar, salix matsudana × salix alba cv. 1002 The importance of monitoring the morphological (conidial dimensions, DNA fingerprints) and physiological (host range and pathogenicity) attributes of local populations of Melampsora and Marssonina species is discussed.     

Inter‐ and intraspecific variation and genotype × site interaction in Salix Alba L., S. Dasyclados Wimm. and S. viminalis L.

Three test series were planted on sites selected for their contrasting properties to study the genotype × site interaction. Various clones of Salix alba (8 clones in the 1980 series), S. dasyclados (8 clones in 1982 series) and S. viminalis (6, 9, 13 clones in the various series) were included. Growth capacity, survival and infection by rust fungi were analysed. Interspecific variation was seen at some of the sites in the 1982 series but it was pronounced in the 1980 series where 5. viminalis outgrew 5. alba at all sites. Intraspecific variation could be seen for growth capacity, survival and rust resistance. A genotype × site interaction was detected for all variables studied. Some of the interaction, especially at the species level, can be explained by the variation in susceptibility to rust fungi and preference for soil types. In the breeding work with Salix the genotype × site interaction must be considered.     

Cronartium flaccidum sporulates on new Euphrasia species in natural habitats in Finland

Natural sporulation of a pine stem rust, Cronartium flaccidum, was investigated in forested habitats in northern Finland. First, a diffuse sample (100) of seven species of hemiparasitic plants was collected in their natural habitats close to one another. C. flaccidum was found to sporulate on six species. The rust was most common on Melampyrum sylvaticum with 21% of the plants and 2% of the leaves carrying telia. Telia occurred occasionally also on Euphrasia frigida, E. nemorosa, E. stricta, Pedicularis palustris ssp. palustris and Melampyrum pratense (less than 5% of the plants and 0.3% of the leaves). The rust was absent on Rhinanthus minor. Second, a diffuse sample of each of M. sylvaticum, M. pratense, Euphrasia sp. and Rhinanthus sp. was collected from 10 sample plots surrounding the city area. C. flaccidum occurred on 0.29% of the leaves of M. sylvaticum, and 0.02%–0.03% of M. pratense and Euphrasia, but it was lacking from Rhinanthus. The results indicated that C. flaccidum was most common and widespread on M. sylvaticum, but sporulated sporadically also on Euphrasia sp., P. palustris and M. pratense. This is the first report of sporulation of C. flaccidum on Euphrasia frigida and E. nemorosa in their natural habitats. In contrast, a second rust observed in this survey, Coleosporium sp. needle rust, was much more common than C. flaccidum on the investigated plants. In the first site, 43%–54% of plants and 8%–17% of leaves of Melampyrum spp. and 0%–13% of plants and 0%–2% of leaves of Euphrasia spp., Rhinanthus minor and Pedicularis palustris carried uredinia or telia. In the second site, 9%–15% of leaves of Melampyrum spp. and 1%–2% of Euphrasia sp. and Rhinanthus sp. carried rust uredinia and/or telia. The results indicate frequent spore dispersal of both Cronartium and Coleosporium in the area, and versatile rust sporulation on hemiparasitic plant species.     

Increased atmospheric carbon dioxide concentration: effects on eucalypt rust (Puccinia psidii), C:N ratio and essential oils in eucalypt clonal plantlets

Evaluation of impacts of high CO₂ atmospheric concentration is strategically important for the development of adaptation measures and sustainable crop management. The objective of this study was to evaluate the effects of increased atmospheric CO₂ concentration on eucalypt rust (Puccinia psidii), C:N ratio, yield and chemical composition of essential oils and growth of eucalypt clonal plantlets. Two clones with different levels of rust resistance were studied: a Eucalyptus urophylla × E. camaldulensis hybrid (VM 01) and an E. urophylla (clone MN 463). The experiments were performed in open‐top chambers (OTCs) with CO₂ mean concentrations (μmol mol^?1) of 399 (unenclosed control), 412 (OTCs with ambient CO₂ concentration) and 508 (OTCs with high CO₂ concentration) and in closed chambers (CCs) with CO₂ mean concentrations of 390, 405, 520 and 700. Increased atmospheric CO₂ concentrations resulted in a decrease in rust pustules per leaf, uredinia per leaf area, spores per uredinia and area under the disease progress curve in VM 01 (hybrid) clonal plantlets. The disease did not occur in MN 463 clonal plantlets, which demonstrated that high CO₂ concentrations did not change the level of rust resistance. Plant growth of the two clones was stimulated by up to 23% in height and 26% in stem diameter in OTCs and by 18% for both clones in CCs. An increased C:N ratio in leaves, stems and roots was observed only for the VM 01 clonal plantlets. Essential oils produced by VM 01 (2.8 g 100 g^?1) and MN 463 (1.4 g 100 g^?1), as well as the major essential oil compounds (80% 1.8‐cineole for VM 01; 50% 1.8‐cineole and 32% α‐pinene for MN 463), were not altered. In this study, increased concentrations of atmospheric CO₂ favourably impacted eucalypt growth and reduced rust severity, while not influencing the production of essential oils.     

Protocol for screening rust disease resistance in Dalbergia sissoo

A protocol has been developed for inoculation of rust disease in Dalbergia sissoo using urediniospores of Maravilia achroa. This is the first time that disease symptoms caused by M. achroa were artificially produced. The method has been successfully employed for screening 15 clones of D. sissoo against rust disease. Clone S‐167 was the most resistant remaining free from infection. Clones S‐57, S‐106 and S‐124 were also resistant. Clones S‐19, S‐24 and S‐89 were susceptible.     

Variation in susceptibility of six pine species and hybrids to pitch canker caused by Fusarium circinatum

Six pine species or hybrids were tested for susceptibility to pitch canker caused by Fusarium  circinatum. Pinus  densiflora, Pinus  thunbergii, Pinus  x rigitaeda (Pinus  rigida × Pinus  taeda), P. rigida × P. x rigitaeda, Pinus  echinata and Pinus  virginiana were inoculated with three spore loads (50, 500 and 5000 per tree) of F. circinatum. External symptoms, lesion length, and the frequency of reisolation of the fungus were investigated. External symptoms were greatest in P. echinata, followed by P. virginiana, however, P. densiflora was not susceptible to F. circinatum. Based on mean lesion lengths, the six pine species or hybrids differed significantly (p < 0.01) in susceptibility to pitch canker. Pinus  echinata sustained the longest lesions, whereas P. densiflora sustained the shortest lesions. The effect of inoculum density was not significant among three spore treatments within species (p = 0.17), although lesion length was slightly greater at higher spore loads over all pine species. The fungus was reisolated from inoculated stems of all pine species tested, even on trees showing little or no damage from the disease. Additional studies are needed to further explore the basis for resistance to pitch canker.     

Fruiting and sporulation of Thekopsora and Chrysomyxa cone rusts in Picea cones and Prunus leaves

Natural fruiting and sporulation of cone rusts were investigated in cones of Picea spp. and leaves of Prunus spp. in a botanical garden in northern Finland in 2007–2012. Thekopsora areolata was the most frequent cone rust in Picea abies cones, where it colonizes the host tissues and hinders normal seed development. Aecia of T. areolata were also common in cones of Picea engelmannii and occasionally in cones of P. glauca. Aecia of T. areolata sporulated in cones that were at least one year old. Chrysomyxa pirolata, another pathogenic cone rust, fruited and sporulated annually but infrequently in current‐year cones of P. abies. The spruce needle rust, Chrysomyxa ledi, fruited and sporulated commonly in current‐year cone scales of P. abies, P. omorika and P. glauca, while P. rubens, P. mariana and P. pungens appeared to be resistant during the study period. Chrysomyxa ledi did not affect seed development in infected cones. Uredinia of T. areolata frequently occurred on leaves of 41 Finnish and Russian cultivars, varieties or subspecies of Prunus padus L. ssp. badus and ssp. borealis and Pr. virginiana both in the botanical garden and in the field, while 13 exotic Prunus spp. lacked rust fruitbodies. All the Pr. padus cultivars were highly susceptible to T. areolata, thus, spreading the rust efficiently to surroundings. This is the first report of aecia of T. areolata in cones of P. engelmannii and P. glauca, and those of C. ledi in cones of P. omorika and P. glauca. Molecular identification confirmed the presence of T. areolata and C. pirolata on all hosts, and all samples of C. ledi belonged to the C. ledi‐rhododendri complex.