Halvor Solheim
Research Professor (OAP Agreement)
Abstract
The genus Ceratocystiopsis (Ophiostomatales, Ascomycota) includes 21 species, which can be found mainly in association with bark beetles in the Northern Hemisphere. A survey of Ceratocystiopsis species associated with bark beetles infesting Picea abies and Pinus sylvestris in Norway yielded 126 isolates, representing Ceratocystiopsis neglecta and Ceratocystiopsis rollhanseniana, and four species described herein as Ceratocystiopsis chalcographii, Ceratocystiopsis debeeria, Ceratocystiopsis norroenii and Ceratocystiopsis troendelagii. The new taxa were morphologically characterised and phylogenetically analysed on the basis of sequence data of multiple loci (ITS, LSU, beta-tubulin (TUB2), calmodulin (CAL) and translation elongation factor 1-alpha (TEF1) genes). Ceratocystiopsis norroenii and C. rollhanseniana were the most frequently isolated species, and the latter species had the wider vector range.
Authors
Y. P. Tan S. L. Bishop-Hurley R. G. Shivas D.A. Cowan G. Maggs-Kölling Sajeewa S.N. Maharachchikumbura U. Pinruan K.L. Bransgrove S. De la Peña-Lastra E. Larsson T. Lebel S. Mahadevakumar A. Mateos E. R. Osieck A. Rigueiro-Rodríguez S. Sommai K. Ajithkumar A. Akulov F. E. Anderson F. Arenas S. Balashov Á. Bañares D.K. Berger M.V. Bianchinotti S. Bien P. Bilański A. G. Boxshall M. Bradshaw J. Broadbridge F.J.S. Calaça C. Campos-Quiroz J. Carrasco-Fernández J.F. Castro S. Chaimongkol S. Chandranayaka Y. Chen D. Comben J.D.W. Dearnaley A.S. Ferreira-Sá K. Dhileepan M.L. Diaz P.K. Divakar S. Xavier-Santos A. Fernández-Bravo J. Gené F.E. Guard M. Guerra S. Gunaseelan J. Houbraken K. Janik-Superson R. Jankowiak M. Jeppson Ž. Jurjević M. Kaliyaperumal L.A. Kelly K. Kezo A.N. Khalid P. Khamsuntorn D. Kidanemariam M. Kiran E. Lacey G.J. Langer L.V. López-Llorca J. J. Luangsa-ard P. Lueangjaroenkit H.T. Lumbsch J. G. Maciá-Vicente L.S. Mamatha Bhanu T.S. Marney J.E. Marqués-Gálvez A. Morte A. Naseer A. Navarro-Ródenas O. Oyedele S. Peters S. Piskorski L. Quijada G.H. Ramírez K. Raja A. Razzaq V.J. Rico A. Rodríguez M. Ruszkiewicz-Michalska R.M. Sánchez C. Santelices A.S. Savitha M. Serrano L. Leonardo-Silva Halvor Solheim S. Somrithipol M.Y. Sreenivasa H. Stępniewska D. Strapagiel T. Taylor D. Torres-Garcia J. Vauras M. Villarreal C.M Visagie M. Wołkowycki W. Yingkunchao E. Zapora J.Z. Groenewald P.W. CrousAbstract
Novel species of fungi described in this study include those from various countries as follows: Argentina, Colletotrichum araujiae on leaves, stems and fruits of Araujia hortorum. Australia, Agaricus pateritonsus on soil, Curvularia fraserae on dying leaf of Bothriochloa insculpta, Curvularia millisiae from yellowing leaf tips of Cyperus aromaticus, Marasmius brunneolorobustus on well-rotted wood, Nigrospora cooperae from necrotic leaf of Heteropogon contortus, Penicillium tealii from the body of a dead spider, Pseudocercospora robertsiorum from leaf spots of Senna tora, Talaromyces atkinsoniae from gills of Marasmius crinis-equi and Zasmidium pearceae from leaf spots of Smilax glyciphylla. Brazil, Preussia bezerrensis fromair. Chile, Paraconiothyrium kelleni from the rhizosphere of Fragaria chiloensis subsp. chiloensis f. chiloensis. Finland, Inocybe udicola onsoilinmixedforest with Betula pendula, Populus tremula, Picea abies and Alnus incana. France, Myrmecridium normannianum on dead culm of unidentified Poaceae. Germany, Vexillomyces fraxinicola from symptomless stem wood of Fraxinus excelsior. India, Diaporthe limoniae on infected fruit of Limonia acidissima, Didymella naikii on leaves of Cajanus cajan, and Fulvifomes mangroviensis on basal trunk of Aegiceras corniculatum. Indonesia, Penicillium ezekielii from Zea mays kernels. Namibia, Neocamarosporium calicoremae and Neocladosporium calicoremae on stems of Calicorema capitata, and Pleiochaeta adenolobi on symptomatic leaves of Adenolobus pechuelii. Netherlands, Chalara pteridii on stems of Pteridium aquilinum, Neomackenziella juncicola (incl. Neomackenziella gen. nov.)and Sporidesmiella junci from dead culms of Juncus effusus. Pakistan, Inocybe longistipitata on soil in a Quercus forest. Poland, Phytophthora viadrina from rhizosphere soil of Quercus robur, and Septoria krystynae on leaf spots of Viscum album. Portugal (Azores), Acrogenospora stellata on dead wood or bark. South Africa, Phyllactinia greyiae on leaves of Greyia sutherlandii and Punctelia anae on bark of Vachellia karroo. Spain, Anteaglonium lusitanicum on decaying wood of Prunus lusitanica subsp. lusitanica, Hawksworthiomyces riparius from fluvial sediments, Lophiostoma carabassense endophytic in roots of Limbarda crithmoides, and Tuber mohedanoi from calcareussoils. Spain (Canary Islands), Mycena laurisilvae on stumps and woody debris. Sweden, Elaphomyces geminus from soil under Quercus robur. Thailand, Lactifluus chiangraiensis on soil under Pinus merkusii, Lactifluus nakhonphanomensis and Xerocomus sisongkhramensis on soil under Dipterocarpus trees. Ukraine, Valsonectria robiniae on dead twigs of Robinia hispida. USA, Spiralomyces americanus (incl. Spiralomyces gen. nov.) from office air. Morphological and culture characteristics are supported by DNA barcodes.
Authors
Ari Hietala Ahto Agan Nina Elisabeth Nagy Isabella Børja Volkmar Timmermann Rein Drenkhan Halvor SolheimAbstract
The populations of European ash and its harmless fungal associate Hymenoscyphus albidus are in decline owing to ash dieback caused by the invasive Hymenoscyphus fraxineus, a fungus that in its native range in Asia is a harmless leaf endophyte of local ash species. To clarify the behavior of H. albidus and its spatial and temporal niche overlap with the invasive relative, we used light microscopy, fungal species-specific qPCR assays, and PacBio long-read amplicon sequencing of the ITS1-5.8S-ITS2 region to examine fungal growth and species composition in attached leaves of European ash. The plant material was collected from a healthy stand in central Norway, where ash saplings in late autumn showed leaflet vein necrosis like that commonly related to H. fraxineus. For reference, leaflet samples were analyzed from stands with epidemic level of ash dieback in southeastern Norway and Estonia. While H. albidus was predominant in the necrotic veins in the healthy stand, H. fraxineus was predominant in the diseased stands. Otherwise, endophytes with pathogenic potential in the genera Venturia (anamorph Fusicladium), Mycosphaerella (anamorph Ramularia), and Phoma, and basidiomycetous yeasts formed the core leaflet mycobiome both in the healthy and diseased stands. In necrotic leaf areas with high levels of either H. albidus or H. fraxineus DNA, one common feature was the high colonization of sclerenchyma and phloem, a region from which the ascomata of both species arise. Our data suggest that H. albidus can induce necrosis in ash leaves, but that owing to low infection pressure, this first takes place in tissues weakened by autumn senescence, 1–2 months later in the season than what is characteristic of H. fraxineus at an epidemic phase of ash dieback. The most striking difference between these fungi would appear to be the high fecundity of H. fraxineus. The adaptation to a host that is phylogenetically closely related to European ash, a tree species with high occurrence frequency in Europe, and the presence of environmental conditions favorable to H. fraxineus life cycle completion in most years may enable the build-up of high infection pressure and challenge of leaf defense prior to autumn senescence.
Abstract
No abstract has been registered
Authors
Kalev Adamson Marili Laas Kathrin Blumenstein Johanna Busskamp Gitta J. Langer Darta Klavina Anu Kaur Tiit Maaten Martin S. Mullett Michael M. Müller Emília Ondrušková Allar Padari Enn Pilt Taavi Riit Halvor Solheim Liina Soonvald Leho Tedersoo Eeva Terhonen Rein DrenkhanAbstract
Diplodia sapinea is a cosmopolitan endophyte and opportunistic pathogen having occurred on several conifer species in Europe for at least 200 years. In Europe, disease outbreaks have increased on several Pinus spp. in the last few decades. In this study, the genetic structure of the European and western Asian D. sapinea population were investigated using 13 microsatellite markers. In total, 425 isolates from 15 countries were analysed. A high clonal fraction and low genetic distance between most subpopulations was found. One single haplotype dominates the European population, being represented by 45.3% of all isolates and found in nearly all investigated countries. Three genetically distinct subpopulations were found: Central/North European, Italian and Georgian. The recently detected subpopulations of D. sapinea in northern Europe (Estonia) share several haplotypes with the German subpopulation. The northern European subpopulations (Latvia, Estonia and Finland) show relatively high genetic diversity compared to those in central Europe suggesting either that the fungus has existed in the North in an asymptomatic/endophytic mode for a long time or that it has spread recently by multiple introductions. Considerable genetic diversity was found even among isolates of a single tree as 16 isolates from a single tree resulted in lower clonal fraction index than most subpopulations in Europe, which might reflect cryptic sexual proliferation. According to currently published allelic patterns, D. sapinea most likely originates from North America or from some unsampled population in Asia or central America. In order to enable the detection of endophytic or latent infections of planting stock by D. sapinea, new species-specific PCR primers (DiSapi-F and Diplo-R) were designed. During the search for Diplodia isolates across the world for species specific primer development, we identified D. africana in California, USA, and in the Canary Islands, which are the first records of this species in North America and in Spain.
Abstract
The relationship between the ecological success of needle pathogens of forest trees and species richness of co-inhabiting endophytic fungi is poorly understood. One of the most dangerous foliar pathogens of pine is Dothistroma septosporum, which is a widely spread threat to northern European forests. We sampled two Pinus sylvestris sites in Estonia and two in Norway in order to analyse the relations between the abundance of D. septosporum and overall fungal richness, specific fungal species composition, time of season, needle age and position in the canopy. In both countries, the overall species richness of fungi was highest in autumn, showing a trend of increase with needle age. The overall species richness in the second-year needles in Estonia and third-year needles in Norway was similar, suggesting that a critical colonization threshold for needle shed in P. sylvestris is breached earlier in Estonia than in Norway. The fungal species richness in P. sylvestris needles was largely affected by Lophodermium conigenum. Especially in older needles, the relative abundance of L. conigenum was significantly higher in spring compared to summer or autumn. The timing of recruitment and colonization mechanisms of different foliage endophytes are shortly discussed.
Abstract
Sporothrix (Sordariales, Ascomycota) is a well-supported monophyletic lineage within the Ophiostomatales, species of which occur in a diverse range of habitats including on forest trees, in the soil, associated with bark beetles and mites as well as on the fruiting bodies of some Basidiomycota. Several species have also been reported as important human and animal pathogens. During surveys of insect- and wound-associated Ophiostomatales from hardwood trees in Poland, many isolates with affinity to Sporothrix were recovered. In the present study, six undescribed Sporothrix spp. collected during these surveys are characterized based on their morphological characteristics and multi-locus phylogenenetic inference. They are described as Sporothrix cavum, Sporothrix cracoviensis, S. cryptarchum, S. fraxini, S. resoviensis, and S. undulata. Two of the Sporothrix spp. reside in the S. gossypina-complex, while one forms part of the S. stenoceras-complex. One Sporothrix sp. is a member of lineage F, and two other species grouped outside any of the currently defined species complexes. All the newly described species were recovered from hardwood habitats in association with sub-cortical insects, wounds or woodpecker cavities. These species were morphologically similar, with predominantly asexual states having hyaline or lightly pigmented conidia, which produce holoblastically on denticulate conidiogenous cells. Five of the new taxa produce ascomata with necks terminating in long ostiolar hyphae and allantoid ascospores without sheaths. The results suggest that Sporothrix species are common members of the Ophiostomatales in hardwood ecosystems of Poland.
Abstract
The scope of this study was to provide an update on fluoride (F) emission effects on vegetation around three aluminium smelters. We visited Årdal and Sunndal smelters in 2019-2020 and Mosjøen in 2020, assessed and documented the visual symptoms of F-damage on vegetation and related these to detected values of F in plant tissue. Three plant species showed qualities as useful indicators: Rowan, pine and St. John’s wort. Because male-fern accumulated extreme F-values and showed clear grazing damage, the monitoring of this species may be warranted because of the potential health hazard for the grazing animals. In Årdal and Sunndal, during 2019 and 2020, we detected the highest F-values in male-fern, ranging from 94 to 925 mg F/kg. In rowan, the highest F-concentration was detected in trees growing within the Årdal smelter (1161 mg F/kg) but on all other locations the F-concentrations in rowan ranged from 4 to 327 mg F/kg. In pine, the F-concentrations ranged from 6-351 mg F/kg for all needle ages, but older needles always accumulated more F than younger ones. In St. John’s wort the accumulated F-values ranged from 10-84 mg F/kg. At all smelters there was a gradient of decreasing F-concentration in vegetation with increasing distance from the smelter. F-emissions in Årdal (12 and 11 kg F/hour in 2019 and 2020, respectively) and in Sunndal during 2019 (12 kg F/hour) were only slightly higher than the recommended limits (10 kg F/hour) for damage on vegetation, while in Mosjøen the F-emissions were 7 kg F/hour in 2020. The presence of F-damage on vegetation was consistent with the reported emission-levels. On basis of this evaluation, reductions in emissions are still advisable in Årdal and Sunndal, while the situation is acceptable in Mosjøen.
Authors
Martin S. Mullett Rein Drenkhan Kalev Adamson Piotr Boroń Anna Lenart-Boroń Irene Barnes Michal Tomšovský Zuzana Jánošíková Katarína Adamčíková Emília Ondrušková Valentin Queloz Barbara Piskur Dmitry L. Musolin Kateryna Davydenko Margarita Georgieva Sophie Schmitz Audrius Kačergius Luisa Ghelardini Jelena Kranjec Orlović Michael Müller Funda Oskay Tine Hauptman Ágnes Halász Svetlana Markovskaja Halvor Solheim Martti Vuorinen Renate Heinzelmann Richard C. Hamelin Adam KonečnýAbstract
Dothistroma septosporum, the primary causal agent of Dothistroma needle blight, is one of the most significant foliar pathogens of pine worldwide. Its wide host and environmental ranges have led to its global success as a pathogen and severe economic damage to pine forests in many regions. This comprehensive global population study elucidated the historical migration pathways of the pathogen to reveal the Eurasian origin of the fungus. When over 3800 isolates were examined, three major population clusters were revealed: North America, Western Europe, and Eastern Europe, with distinct subclusters in the highly diverse Eastern European cluster. Modeling of historical scenarios using approximate Bayesian computation revealed the North American cluster was derived from an ancestral population in Eurasia. The Northeastern European subcluster was shown to be ancestral to all other European clusters and subclusters. The Turkish subcluster diverged first, followed by the Central European subcluster, then the Western European cluster, which has subsequently spread to much of the Southern Hemisphere. All clusters and subclusters contained both mating-types of the fungus, indicating the potential for sexual reproduction, although asexual reproduction remained the primary mode of reproduction. The study strongly suggests the native range of D. septosporum to be in Eastern Europe (i.e., the Baltic and Western Russia) and Western Asia.
Authors
Hanno Sandvik Olga Hilmo Snorre Henriksen Reidar Elven Per Arvid Åsen Hanne Hegre Oddvar Pedersen Per Anker Pedersen Heidi Solstad Vigdis Vandvik Kristine Bakke Westergaard Frode Ødegaard Sandra Charlotte Helene Åström Hallvard Elven Anders Endrestøl Øivind Gammelmo Bjørn Arild Hatteland Halvor Solheim Björn Nordén Leif Sundheim Venche Talgø Tone Falkenhaug Bjørn Gulliksen Anders Jelmert Eivind Oug Jan Henry Sundet Elisabet Forsgren Anders Gravbrøt Finstad Trygve H. Hesthagen Kjell Harald Nedreaas Rupert Wienerroither Vivian Husa Stein Fredriksen Kjersti Sjøtun Henning Steen Haakon Hansen Inger Sofie Hamnes Egil Karlsbakk Christer Magnusson Bjørnar Ytrehus Hans Christian Pedersen Jon Swenson Per Ole Syvertsen Bård Gunnar Stokke Jan Ove Gjershaug Dag Dolmen Gaute Kjærstad Stein Ivar Johnsen Thomas Correll Jensen Kristian Hassel Lisbeth GederaasAbstract
1. Due to globalisation, trade and transport, the spread of alien species is increasing dramatically. Some alien species become ecologically harmful by threatening native biota. This can lead to irreversible changes in local biodiversity and ecosystem functioning, and, ultimately, to biotic homogenisation. 2. We risk-assessed all alien plants, animals, fungi and algae, within certain delimitations, that are known to reproduce in Norway. Mainland Norway and the Arctic archipelago of Svalbard plus Jan Mayen were treated as separate assessment areas. Assessments followed the Generic Ecological Impact Assessment of Alien Species (GEIAA) protocol, which uses a fully quantitative set of criteria. 3. A total of 1519 species were risk-assessed, of which 1183 were species reproducing in mainland Norway. Among these, 9% were assessed to have a severe impact, 7% high impact, 7% potentially high impact, and 49% low impact, whereas 29% had no known impact. In Svalbard, 16 alien species were reproducing, one of which with a severe impact. 4. The impact assessments also covered 319 so-called door-knockers, i.e. species that are likely to establish in Norway within 50 years, and 12 regionally alien species. Of the door-knockers, 8% and 10% were assessed to have a severe and high impact, respectively. 5. The impact category of most species was driven by negative interactions with native species, transformation of threatened ecosystems, or genetic contamination. The proportion of alien species with high or severe impact varied significantly across the different pathways of introduction, taxonomic groups, time of introduction, and the environments colonised, but not across continents of origin. 6. Given the large number of alien species reproducing in Norway and the preponderance of species with low impact, it is neither realistic nor necessary to eradicate all of them. Our results can guide management authorities in two ways. First, the use of quantitative assessment criteria facilitates the prioritisation of management resources across species. Second, the background information collected for each species, such as introduction pathways, area of occupancy and ecosystems affected, helps designing appropriate management measures.
Authors
Hanno Sandvik Olga Hilmo Snorre Henriksen Reidar Elven Per Arvid Åsen Hanne Hegre Oddvar Pedersen Per Anker Pedersen Heidi Solstad Vigdis Vandvik Kristine Bakke Westergaard Frode Ødegaard Sandra Charlotte Helene Åström Hallvard Elven Anders Endrestøl Øivind Gammelmo Bjørn Arild Hatteland Halvor Solheim Björn Nordén Leif Sundheim Venche Talgø Tone Falkenhaug Bjørn Gulliksen Anders Jelmert Eivind Oug Jan Henry Sundet Elisabet Forsgren Anders Gravbrøt Finstad Trygve H. Hesthagen Kjell Harald Nedreaas Rupert Wienerroither Vivian Husa Stein Fredriksen Kjersti Sjøtun Henning Steen Haakon Hansen Inger Sofie Hamnes Egil Karlsbakk Christer Magnusson Bjørnar Ytrehus Hans Christian Pedersen Jon Swenson Per Ole Syvertsen Bård Gunnar Stokke Jan Ove Gjershaug Dag Dolmen Gaute Kjærstad Stein Ivar Johnsen Thomas Correll Jensen Kristian Hassel Lisbeth GederaasAbstract
No abstract has been registered
Authors
Melissa Magerøy Erik Christiansen Bo Långström Anna-Karin Borg-Karlson Halvor Solheim Niklas Björklund Tao Zhao Axel Schmidt Carl Gunnar Fossdal Paal KrokeneAbstract
Plants can form an immunological memory known as defense priming, whereby exposure to a priming stimulus enables quicker or stronger response to subsequent attack by pests and pathogens. Such priming of inducible defenses provides increased protection and reduces allocation costs of defense. Defense priming has been widely studied for short‐lived model plants such as Arabidopsis, but little is known about this phenomenon in long‐lived plants like spruce. We compared the effects of pretreatment with sublethal fungal inoculations or application of the phytohormone methyl jasmonate (MeJA) on the resistance of 48‐year‐old Norway spruce (Picea abies) trees to mass attack by a tree‐killing bark beetle beginning 35 days later. Bark beetles heavily infested and killed untreated trees but largely avoided fungus‐inoculated trees and MeJA‐treated trees. Quantification of defensive terpenes at the time of bark beetle attack showed fungal inoculation induced 91‐fold higher terpene concentrations compared with untreated trees, whereas application of MeJA did not significantly increase terpenes. These results indicate that resistance in fungus‐inoculated trees is a result of direct induction of defenses, whereas resistance in MeJA‐treated trees is due to defense priming. This work extends our knowledge of defense priming from model plants to an ecologically important tree species.
Abstract
During surveys of insect-associated mycobiomes in Norway, Poland, and Russia, isolates with affinity to Graphilbum (Ophiostomatales, Ascomycota) were recovered. In this study, eight known Graphilbum species as well as the newly collected isolates were compared based on morphology and DNA sequence data for four gene regions. The results revealed seven new species, described here as G. acuminatum, G. carpaticum, G. curvidentis, G. furuicola, G. gorcense, G. interstitiale, and G. sexdentatum. In addition to these species, G. crescericum and G. sparsum were commonly found in Norway. All new species were recovered from conifers in association with bark beetles, cerambycid beetles, and weevils and were morphologically similar, predominantly with pesotum-like asexual morphs. Where sexual morphs were present, these were small ascomata with short necks and rodshaped ascospores having hyaline sheaths. The results suggest that Graphilbum species are common members of the Ophiostomatales in conifer ecosystems.
Authors
Ahto Agan Rein Drenkhan Kalev Adamson Leho Tedersoo Halvor Solheim Isabella Børja Iryna Matsiakh Volkmar Timmermann Nina Elisabeth Nagy Ari HietalaAbstract
European ash (Fraxinus excelsior) is threatened by the invasive ascomycete Hymenoscyphus fraxineus originating from Asia. Ash leaf tissues serve as a route for shoot infection but also as a sporulation substrate for this pathogen. Knowledge of the leaf niche partitioning by indigenous fungi and H. fraxineus is needed to understand the fungal community receptiveness to the invasion. We subjected DNA extracted from unwashed and washed leaflets of healthy and diseased European ash to PacBio sequencing of the fungal ITS1-5.8S-ITS2 rDNA region. Leaflets from co-inhabiting rowan trees (Sorbus aucuparia) served as a reference. The overlap in leaflet mycobiomes between ash and rowan was remarkably high, but unlike in rowan, in ash leaflets the sequence read proportion, and the qPCR-based DNA amount estimates of H. fraxineus increased vigorously towards autumn, concomitant with a significant decline in overall fungal richness. The niche of ash and rowan leaves was dominated by epiphytic propagules (Vishniacozyma yeasts, the dimorphic fungus Aureobasidion pullulans and the dematiaceous hyphomycete Cladosporium ramotenellum and H. fraxineus), and endophytic thalli of biotrophs (Phyllactinia and Taphrina species), the indigenous necrotroph Venturia fraxini and H. fraxineus. Mycobiome comparison between healthy and symptomatic European ash leaflets revealed no significant differences in relative abundance of H. fraxineus, but A. pullulans was more prevalent in symptomatic trees. The impacts of host specificity, spatiotemporal niche partitioning, species carbon utilization profiles and life cycle traits are discussed to understand the ecological success of H. fraxineus in Europe. Further, the inherent limitations of different experimental approaches in the profiling of foliicolous fungi are addressed.
Authors
Beata Strzałka Robert Jankowiak Piotr Bilański Nikita Patel Georg Hausner Riikka Linnakoski Halvor SolheimAbstract
Bark beetles belonging to the genus Dryocoetes (Coleoptera, Curculionidae, Scolytinae) are known vectors of fungi, such as the pathogenic species Grosmannia dryocoetidis involved in alpine fir (Abies lasiocarpa) mortality. Associations between hardwood-infesting Dryocoetes species and fungi in Europe have received very little research attention. Ectosymbiotic fungi residing in Ceratocystiopsis and Leptographium (Ophiostomatales, Sordariomycetes, Ascomycota) were commonly detected in previous surveys of the Dryocoetes alni-associated mycobiome in Poland. The aim of this study was to accurately identify these isolates and to provide descriptions of the new species. The identification was conducted based on morphology and DNA sequence data for six loci (ITS1-5.8S, ITS2-28S, ACT, CAL, TUB2, and TEF1-α). This revealed two new species, described here as Ceratocystiopsis synnemata sp. nov. and Leptographium alneum sp. nov. The host trees for the new species included Alnus incana and Populus tremula. Ceratocystiopsis synnemata can be distinguished from its closely related species, C. pallidobrunnea, based on conidia morphology and conidiophores that aggregate in loosely arranged synnemata. Leptographium alneum is closely related to Grosmannia crassivaginata and differs from this species in having a larger ascomatal neck, and the presence of larger club-shaped cells.
Authors
Hanno Sandvik Dag Dolmen Reidar Elven Tone Falkenhaug Elisabet Forsgren Haakon Hansen Kristian Hassel Vivian Husa Gaute Kjærstad Frode Ødegaard Hans Christian Pedersen Halvor Solheim Bård Gunnar Stokke Per Arvid Åsen Sandra Åström Tor Erik Brandrud Hallvard Elven Anders Endrestøl Anders Gravbrøt Finstad Stein Fredriksen Øivind Gammelmo Jan Ove Gjershaug Bjørn Gulliksen Inger Hamnes Bjørn Arild Hatteland Hanne Hegre Trygve Hesthagen Anders Jelmert Thomas C Jensen Stein Ivar Johnsen Egil Karlsbakk Christer Magnusson Kjell Nedreaas Björn Nordén Eivind Oug Oddvar Pedersen Per Anker Pedersen Kjersti Sjøtun Jon Kristian Skei Heidi Solstad Leif Sundheim Jon E Swenson Per Ole Syvertsen Venche Talgø Vigdis Vandvik Kristine B Westergaard Rupert Wienerroither Bjørnar Ytrehus Olga Hilmo Snorre Henriksen Lisbeth GederaasAbstract
We present the results of an inventory and status assessment of alien species in Norway. The inventory covered all known multicellular neobiota, 2496 in total, 1039 of which were classified as naturalised. The latter constitute c. 3% of all species known to be stably reproducing in Norway. These figures are higher than expected from Norway’s latitude, which may be due a combination of climatic and historical factors, as well as sampling effort. Most of the naturalised neobiota were plants (71%),followed by animals (21%) and fungi (8%). The main habitat types colonised were open lowlands (79%), urban environments (52%) and woodlands (42%). The main areas of origin were Europe (67%), North America (15%) and Asia (13%). For most taxa, the rate of novel introductions seems to have been increasing during recent decades. Within Norway, the number of alien species recorded per county was negatively correlated with latitude and positively correlated with human population density. In the high-Arctic territories under Norwegian sovereignty, i.e. Svalbard and Jan Mayen, 104 alien species were recorded, of which 5 were naturalised.
Authors
R. Ioos P. Chrétien J. Perrault C. Jeandel C. Dutech P. Gonthier F. Sillo Ari Hietala Halvor Solheim J. HubertAbstract
Four species of the destructive forest pathogen Heterobasidion annosum sensu lato (s.l.) are present in Europe: H. annosum sensu stricto (s.s.), H. abietinum and H. parviporum are native species, while H. irregulare is a non‐native invasive species currently reported only in Italy, yet recommended for regulation throughout Europe. In this study, real‐time PCR detection tests were developed for each of the four species, which can be used simultaneously or individually thanks to probes labelled with species‐specific fluorescent dyes. The different performance criteria of each assay were evaluated, and it was determined that they were theoretically capable of detecting amounts of DNA corresponding to 311, 29 and 29 cell nuclei in H. annosum s.s., H. irregulare and H. parviporum, respectively. The specificity of each assay was assessed with a wide set of strains. Real‐time PCR tests successfully detected Heterobasidion species from 36 fruiting bodies taken from the forest, as well as from artificially inoculated or naturally infected wood samples. The multiplex real‐time PCR assays developed in this study could have practical applications both in forest management and in phytosanitary monitoring.
Abstract
No abstract has been registered
Authors
Leif Sundheim Christer Magnusson Arild Sletten Per Hans Micael Wendell Guro Brodal Åshild Ergon Halvor Solheim Anne Marte Tronsmo Trond RafossAbstract
No abstract has been registered
Authors
Ari Hietala Isabella Børja Hugh B. Cross Nina Elisabeth Nagy Jørn Henrik Sønstebø Volkmar Timmermann Adam Vivian-Smith Halvor SolheimAbstract
No abstract has been registered
Abstract
No abstract has been registered
Authors
Ari Hietala Isabella Børja Hugh Cross Nina Elisabeth Nagy Halvor Solheim Volkmar Timmermann Adam Vivian-SmithAbstract
European ash (Fraxinus excelsior), a keystone species with wide distribution and habitat range in Europe, is threatened at a continental scale by an invasive alien ascomycete, Hymenoscyphus fraxineus. In its native range of Asia, this fungus is a leaf endophyte with weak parasitic capacity and robust saprobic competence in local ash species that are closely related to European ash. In European ash, H. fraxineus has a similar functional role as in Asia, but the fungus also aggressively kills shoots, resulting in crown dieback and tree death. H. fraxineus is a typical invasive species, as its spread relies on high propagule pressure. While crown dieback of European ash is the most obvious symptom of ash dieback, the annual colonization of ash leaves is a crucial key dependency for the invasiveness of H. fraxineus, since its fruiting bodies are formed on overwintered leaf vein tissues in soil debris. Leaves of European ash host a wide range of indigenous epiphytes, endophytes, facultative parasites and biotrophic fungi, including Hymenoscyphus albidus, a relative of H. fraxineus that competes for the same sporulation niche as the invader. At face value, leaves of European ash are colonized by a large and diverse indigenous mycobiome. In order to understand why this invader became successful in Europe, we discuss and summarize the current knowledge of diversity, seasonal dynamics and traits of H. fraxineus and indigenous fungi associated with leaves of European ash.
Abstract
Ophiostoma spp. (Ophiostomatales, Ascomycota) are well-known fungi associated with bark and ambrosia beetles (Curculionidae: Scolytinae, Platypodinae). Fungi in the Ophiostomatales include serious tree pathogens as well as agents of timber blue-stain. Although these fungi have been extensively studied in the northern hemisphere, very little is known regarding their occurrence on hardwoods in Europe. The aims of the present study were to identify and characterize new Ophiostoma spp. associated with bark and ambrosia beetles infesting hardwoods in Norway and Poland, and to resolve phylogenetic relationships of Ophiostoma spp. related to the Norwegian and Polish isolates, using multigene phylogenetic analyses. Results obtained from five gene regions (ITS, LSU, b-tubulin, calmodulin, translation elongation factor 1-a) revealed four new Ophiostoma spp. These include Ophiostoma hylesinum sp. nov., O. signatum sp. nov., and O. villosum sp. nov. that phylogenetically are positioned within the Ophiostoma ulmi complex. The other new species, Ophiostoma pseudokarelicum sp. nov. reside along with Ophiostoma karelicum in a discrete, well-supported phylogenetic group in Ophiostoma s. stricto. The results of this study clearly show that the diversity and ecology of Ophiostoma spp. on hardwoods in Europe is poorly understood and that further studies are required to enrich our knowledge about these fungi.
Authors
Isabella Børja Kjell Andreassen Jan Čermák Lise Dalsgaard Arthur Gessler Douglas Lawrence Godbold Rainer Hentschel Zachary E. Kayler Paal Krokene Nadezhda Nadezhdina Sabine Rosner Halvor Solheim Jan Svetlik Mari Mette Tollefsrud Ole Einar TveitoAbstract
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Authors
Louise Eriksson Johanna Boberg Thomas L. Cech Tamara Corcobado Marie-Laure Desprez-Loustau Ari Hietala Marilia Horta Jung Thomas Jung Hatice Tugba Dogmus Lehtijarvi Funda Oskay Slavtcho Slavov Halvor Solheim Jan Stenlid Jonas OlivaAbstract
Political action can reduce introductions of diseases caused by invasive forest pathogens (IPs) and public support is important for effective prevention. The public’s awareness of IP problems and the acceptability of policies aiming to combat these pathogens were surveyed in nine European countries (N = 3469). Although awareness of specific diseases (e.g., ash dieback) varied, problem awareness and policy acceptability were similar across countries. The public was positive towards policies for informational measures and stricter standards for plant production, but less positive towards restricting public access to protected areas. Multilevel models, including individual and country level variables, revealed that media exposure was positively associated with awareness of IP problems, and strengthened the link between problem awareness and policy acceptability. Results suggest that learning about IPs through the media and recognizing the associated problems increase policy acceptability. Overall, the study elaborates on the anthropogenic dimension of diseases caused by IPs.
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Dieback of European ash, caused by the ascomycete Hymenoscyphus fraxineus originating from Asia, has rapidly spread across Europe, and is threatening this keystone tree at a continental scale. High propagule pressure is characteristic to invasive species. Consistently, the enormous production of windborne ascospores by H. fraxineus in an ash forest with epidemic level of disease obviously facilitates its invasiveness and long distance spread. To understand the rate of build-up of propagule pressure by this pathogen following its local introduction, during 2011–2017 we monitored its sporulation at a newly infested ash stand in south-western Norway characterized with mild winters and cool summers. We also monitored the propagule pressure by Hymenoscyphus albidus, a non-pathogenic native species that competes for the same sporulation niche with H. fraxineus. During the monitoring period, crown condition of ash trees had impaired, and 20% of the dominant trees were severely damaged in 2017. H. fraxineus showed an exponential increase in spore production between 2012 and 2015, followed by drastic decline in 2016 and 2017. During 2011–2013, the two Hymenoscyphus species showed similar sporulation level, but thereafter spores of H. albidus were no longer detected. The data suggest that following local introduction, the population of H. fraxineus reaches rapidly an exponential growth stage if the local weather conditions are favorable for ascomata maturation across years. In the North Atlantic climate, summer temperatures critically influence the pathogen infection pressure, warm summers allowing the population to grow according to its biotic potential, whereas cold summers can cause a drastic decline in propagule pressure.
Authors
Mark McMullan Maryam Rafiqi Gemy Kaithakottil Bernardo J. Clavijo Lorelei Bilham Elizabeth Orton Lawrence Percival-Alwyn Ben J. Ward Anne Edwards Diane G.O. Saunders Gonzalo Garcia Accinelli Jonathan Wright Walter Verweij Georgios Koutsovoulos Kentaro Yoshida Tsuyoshi Hosoya Louisa Williamson Philip Jennings Renaud Ioos Claude Husson Ari Hietala Adam Vivian-Smith Halvor Solheim Dan MaClean Christine Fosker Neil Hall James K.M. Brown David Swarbreck Mark Blaxter J. Allan Downie Matthew D. ClarkAbstract
Accelerating international trade and climate change make pathogen spread an increasing concern. Hymenoscyphus fraxineus, the causal agent of ash dieback, is a fungal pathogen that has been moving across continents and hosts from Asian to European ash. Most European common ash trees (Fraxinus excelsior) are highly susceptible to H.fraxineus, although a minority (~5%) have partial resistance to dieback. Here, we assemble and annotate a H.fraxineus draft genome, which approaches chromosome scale. Pathogen genetic diversity across Europe and in Japan, reveals a strong bottleneck in Europe, though a signal of adaptive diversity remains in key host interaction genes. We find that the European population was founded by two divergent haploid individuals. Divergence between these haplotypes represents the ancestral polymorphism within a large source population. Subsequent introduction from this source would greatly increase adaptive potential of the pathogen. Thus, further introgression of H.fraxineus into Europe represents a potential threat and Europe-wide biological security measures are needed to manage this disease.
Authors
Robert Jankowiak Agnieszka Ostafińska Truls Aas Halvor Solheim Piotr Bilański Riikka Linnakoski Georg HausnerAbstract
Species of Leptographium are characterized by mononematous or synnematous conidiophores and are commonly associated with different arthropods. Some of them also produce a sexual state characterised by globose ascomata with elongated necks. Compared to investigations on coniferous trees, the occurrence of Leptographium species on hardwood trees has been poorly studied in Europe. During a survey of ophiostomatoid fungi on various hardwood tree species in Norway and Poland, three unusual species, which fit in the broader morphological description of Leptographium spp., were found in association with Trypodendron domesticum, Trypodendron signatum and Dryocoetes alni, and from wounds on a variety of hardwoods. Phylogenetic analyses of sequence data for six different loci (ITS1–5.8 S–ITS2, ITS2-LSU, ACT, b-tubulin, CAL, and TEF-1a) showed that these Leptographium species are phylogenetically closely related to the species of the Grosmannia olivacea complex. The first species forms a well-supported lineage that includes Ophiostoma brevicolle, while the two other new taxa resided in a separate lineage; possibly affiliated with Grosmannia francke-grosmanniae. All the new species produce perithecia with necks terminating in ostiolar hyphae and orange-section shaped ascospores with cucullate, gelatinous sheaths. These species also produce dark olivaceous mononematous asexual states in culture. In addition, two of the newly described species have a second type of conidiophore with a short and non-pigmented stipe. The new Leptographium species can be easily distinguished from each other by their appearance and growth in culture. Based on novel morphological characters and distinct DNA sequences, these fungi were recognised as new taxa for which the names Leptographium tardum sp. nov., Leptographium vulnerum sp. nov., and Leptographium flavum sp. nov. are provided.
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Authors
Isabella Børja Volkmar Timmermann Ari Hietala Mari Mette Tollefsrud Nina Elisabeth Nagy Adam Vivian-Smith Hugh Cross Jørn Henrik Sønstebø Tor Myking Halvor SolheimAbstract
In Norway the common ash (Fraxinus excelsior L.) has its northernmost distribution in Europe. It grows along the coastal range as small fragmented populations. The first occurrence of ash dieback caused by Hymenoscyphus fraxineus in Norway was reported in 2008. At that time, the disease had already spread through large areas of southern and south-eastern parts of Norway. Since then the disease continued spreading with a speed of about 50- 60 km per year along the western coastal range. To monitor the disease development over time, we established eight permanent monitoring plots in south-eastern and western Norway in 2009 and 2012, respectively. In all plots tree mortality was high, especially among the youngest trees in south-eastern Norway. The extent of crown damage has continually increased in all diameter classes for both regions. In 2009, 76.8 % of all trees on the five monitoring plots in south-eastern Norway were considered to be healthy or slightly damaged, and only 8.9 % to be severely damaged. In 2015, 51.7 % were dead, 13.5 % severely damaged and only 25.7 % remained healthy or slightly damaged. To assess the infection pressure and spore dispersal patterns of the pathogen, we used a Burkard volumetric spore sampler placed in an infested ash stand in southern Norway. We examined the airborne ascospores of H. fraxineus and H. albidus captured on the sampling tape microscopically and with real-time PCR assays specific to these fungi. We detected very few ascospores of H. albidus, whereas ascospores of H. fraxineus dominated throughout entire sampling periods of 2009, 2010 and 2011. Spore discharge occurred mainly between the hours of 5 and 8 a.m., though the distinctive sporulation had yearly variation between 5-7 a.m. We observed the same diurnal pattern throughout the entire sampling period, with a seasonal peak in spore liberation between mid-July and midAugust, after which the number of ascospores decreased substantially. Similar diurnal patterns were observed throughout the sampling period except that after mid-August the number of trapped ascospores substantially decreased. To compare the genetic pattern of common ash in the northern and central ranges of Europe we analyzed the Norwegian samples together with available samples from central Europe by using chloroplast and nuclear microsatellite markers. We found that the northern range of common ash was colonized via a single migration route that originated in eastern or south-eastern Europe with little influence originating from other southern or western European refugia. In the northern range margins, genetic diversity decreased and population differentiation increased, coherent with a post-glacial colonization history characterized by founder events and population fluctuations. Based on our findings we discuss the future management and conservational implications.
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Dieback of European ash (Fraxinus excelsior L.), a disease caused by the ascomycete Hymenoscyphus fraxineus (previously referred to as H. pseudoalbidus or Chalara fraxinea), was first observed in Poland in the early 1990ies, and is currently present almost throughout the entire distribution area of European ash. The characteristic symptoms of the disease include dead shoots with necrotic lesions in the bark and discoloration of xylem and pith but the seasonal dynamics of pathogen spread in shoot tissues remain poorly understood. To investigate whether the internal spread of the fungus involves season-specific patterns, saplings with necrotic bark lesions in 1-2 -year-old stem regions were collected during 2014-2015 at time intervals in spring, summer, autumn and winter at several localities in western Ukraine and at two localities in south-eastern Norway. Tissuespecific presence of H. fraxineus was determined by a highly sensitive quantitative real-time PCR assay that is specific to DNA of H. fraxineus. The relatively high proportion of bark samples positive for H. fraxineus in the saplings collected during spring provides support to a model that H. fraxineus can be a primary causative agent of bark lesions and that other fungi may eventually replace it in old infection areas.
Authors
Isabella Børja Kjell Andreassen Jan Čermák Lise Dalsgaard Arthur Gessler Douglas L. Godbold Rainer Hentschel Zachary E. Kayler Paal Krokene Nadezhda Nadezhdina Sabine Rosner Svein Solberg Halvor Solheim Jan Svetlik Mari Mette Tollefsrud Ole Einar TveitoAbstract
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Authors
Ned B. Klopfenstein Jane E. Stewart Yuko Ota John W. Hanna Bryce A. Richardson Amy L. Ross-Davis Rubén D. Elías-Román Kari Korhonen Nenad Djuro Keča Eugenia Iturritxa, Dionicio Alvarado-Rosales Halvor Solheim Nicholas J. Brazee Piotr Łakomy Michelle R. Cleary Eri Hasegawa Taisei Kikuchi Fortunato Garza-Ocañas Panaghiotis Tsopelas Daniel Rigling Simone Prospero, Tetyana Tsykun Jean A. Bérubé Franck O.P. Stefani Saeideh Jafarpour Vladimír Antonín Michal Tomšovský Geral I. McDonald Stephen Woodward Mee-Sook KimAbstract
Armillaria possesses several intriguing characteristics that have inspired wide interest in understanding phylogenetic relationships within and among species of this genus. Nuclear ribosomal DNA sequence– based analyses of Armillaria provide only limited information for phylogenetic studies among widely divergent taxa. More recent studies have shown that translation elongation factor 1-α (tef1) sequences are highly informative for phylogenetic analysis of Armillaria species within diverse global regions. This study used Neighbor-net and coalescence-based Bayesian analyses to examine phylogenetic relationships of newly determined and existing tef1 sequences derived from diverse Armillaria species from across the Northern Hemisphere, with Southern Hemisphere Armillaria species included for reference. Based on the Bayesian analysis of tef1 sequences, Armillaria species from the Northern Hemisphere are generally contained within the following four superclades, which are named according to the specific epithet of the most frequently cited species within the superclade: (i) Socialis/Tabescens (exannulate) superclade including Eurasian A. ectypa, North American A. socialis (A. tabescens), and Eurasian A. socialis (A. tabescens) clades; (ii) Mellea superclade including undescribed annulate North American Armillaria sp. (Mexico) and four separate clades of A. mellea (Europe and Iran, eastern Asia, and two groups from North America); (iii) Gallica superclade including Armillaria Nag E (Japan), multiple clades of A. gallica (Asia and Europe), A. calvescens (eastern North America), A. cepistipes (North America), A. altimontana (western USA), A. nabsnona (North America and Japan), and at least two A. gallica clades (North America); and (iv) Solidipes/Ostoyae superclade including two A. solidipes/ostoyae clades (North America), A. gemina (eastern USA), A. solidipes/ostoyae (Eurasia), A. cepistipes (Europe and Japan), A. sinapina (North America and Japan), and A. borealis (Eurasia) clade 2. Of note is that A. borealis (Eurasia) clade 1 appears basal to the Solidipes/Ostoyae and Gallica superclades. The Neighbor-net analysis showed similar phylogenetic relationships. This study further demonstrates the utility of tef1 for global phylogenetic studies of Armillaria species and provides critical insights into multiple taxonomic issues that warrant further study.
Abstract
The goal of this study was to assess the long-term effects of partial harvesting and supplementary soil scarification on the frequency of root and butt rot in managed uneven-sized Norway spruce stands. Frequency of rot and the population structure of the rot fungi were assessed on 1353 stumps after clear-cutting 21 years after a selection harvesting experiment. The initial experiment was comprised of three harvest strength (low, intermediate and high) of single-tree selection, removing approximately 25, 45 and 65% of the stand basal area. Uncut control plots were established at the same time. Supplementary soil scarification was applied in subplots within the single-tree selection plots, using a medium-sized excavator. After clear-cutting the stumps were analyzed with respect to rot caused by Heterobasidion parviporum, Armillaria spp., Stereum sanguinolentum as well as other rot fungi. Rot caused by Armillaria spp. was most common (8.6% of the stumps), while infection by H. parviporum (2.9%) or S. sanguinolentum (3.0%) was less frequent. The group “other rot” (5.4%) comprised 21 identified taxa, each occurring in 1–15 stumps. Significantly lower rot frequencies were found for the uncut control (16.3%) and intermediate harvest strength (15.7%), compared with low harvest strength (23.6%). A rot frequency of 21.0% was found in the high harvest strength. In two of three harvest strengths, the rot frequency was higher than for the uncut control. As the observed rot frequencies did not increase consistently with increasing harvest strength, the results do not completely support the initial expectations of increased rot after single-tree selection compared with the uncut control. However, since the probability of rot in individual stumps on plots treated with single-tree selection was significantly affected by the distance to the nearest strip road (H. parviporum) as well as dependent on the size of and distance to the nearest stump of trees cut during the experimental harvest (H. parviporum, S. sanguinolentum and total rot), it is evident that the single-tree selection harvesting was partially responsible for some of the observed rot. One of the selection criteria in the initial harvest was a sanitary removal of trees of poor vitality. Varying degrees of sanitation felling may therefore have offset the effects of new infections in wounds or spread of rot fungi through adjacent stumps. Supplementary soil scarification in small gaps of the residual stand had no significant effect on the frequency of rot, suggesting that such treatment may be used to facilitate regeneration in uneven-sized spruce stands on similar sites.
Authors
Kalev Adamson Martin S. Mullett Halvor Solheim Irene Barnes Michael M. Müller Jarkko Hantula Martti Vuorinen Audrius Kačergius Svetlana Markovskaja Dmitry L. Musolin Kateryna Davydenko Nenad Keča Karli Ligi Rasa D. Priedite Hanna Millberg Rein DrenkhanAbstract
Dothistroma septosporum, a notorious pine needle pathogen with an unknown historical geographic origin and poorly known distribution pathways, is nowadays found almost in all areas inhabited by pines (Pinus spp.). The main aim of this study was to determine the relationship between North European and East Asian populations. In total, 238 Eurasian D. septosporum isolates from 11 countries, including 211 isolates from northern Europe, 16 isolates from Russian Far East and 11 isolates from Bhutan were analysed using 11 species-specific microsatellite and mating type markers. The most diverse populations were found in northern Europe, including the Baltic countries, Finland and European Russia. Notably, D. septosporum has not caused heavy damage to P. sylvestris in northern Europe, which may suggest a long co-existence of the host and the pathogen. No indication was obtained that the Russian Far East or Bhutan could be the indigenous area of D. septosporum, as the genetic diversity of the fungus there was low and evidence suggests gene flow from northern Europe to Russian Far East. On the western coast of Norway, a unique genetic pattern was observed, which differed from haplotypes dominating other Fennoscandian populations. As an agent of dothistroma needle blight, only D. septosporum was documented in northern Europe and Asia, while D. pini was found in Ukraine and Serbia.
Authors
Kalev Adamson Rein Drenkhan Martin Mullett Halvor Solheim Irene Barnes Michael Müller Jarkko HantulaAbstract
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Ash dieback, caused by the ascomycete Hymenoscyphus fraxineus, has been spreading throughout Europe since the early 1990s, threatening European ash at a continental scale. Little is known about the development of the disease in individual forest trees and in different age classes. In this study we monitored ash dieback on trees of different diameter classes in five permanent plots in ash stands in south-eastern Norway from 2009 to 2016, and from 2012 to 2016 in three plots in western Norway with a shorter disease history. Our results showed that more than 80% of the youngest and more than 40% of the intermediate future crop trees in the plots in south-eastern Norway were dead by 2016, while the disease development in large, dominant trees was slower. Although less damage has been observed in the plots in western Norway, the trend for the juvenile trees is the same as in south-eastern Norway with rapidly increasing damage and mortality. Most dead trees in south-eastern Norway were found at sites with high soil moisture and showed symptoms of root-rot caused by Armillaria species. Infected trees, both young and old ones, are weakened by the disease and appear to be more susceptible to other, secondary pathogens, especially under unfavourable site conditions.
Authors
Leif Sundheim Daniel Flø Micael Wendell Guro Brodal Åshild Ergon Christer Magnusson Arild Sletten Halvor Solheim Anne Marte Tronsmo Bjørn Økland Trond RafossAbstract
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Abstract
Ash dieback, caused by the ascomycete Hymenoscyphus fraxineus, was first observed in the eastern and southernmost Norway in 2008. Based on the age of stem bark lesions, it was concluded that the fungus had arrived to the region no later than 2006. Since 2008 the annual spread of the disease northwards along the west coast of Norway has been monitored. The registration was done each year during early summer around a disease frontier recorded in the previous year. The occurrence of necrotic bark lesions in the previous-year shoots and dieback of these shoots, and isolation of H. fraxineus from the discoloured wood associated with necrotic bark lesions were used as signs of ash dieback. These records indicate an annual spread of ash dieback in the range between 25 km and 78 km, and a mean annual spread of 51 km. The cause of the spread is discussed.
Authors
Robert Jankowiak Beata Strzałka Piotr Bilański Riikka Linnakoski Truls Aas Halvor Solheim Małgorzata Groszek Z. Wilhelm de BeerAbstract
Species of Leptographium are generally characterized by mononematous conidiophores and are commonly associated with bark beetles and weevils. These species are responsible for sapstain and in some cases serious diseases on a range of primarily coniferous trees. In comparison with coniferous trees, the occurrence of Leptographium species on hardwood trees has been poorly studied in Europe. During a survey of ophiostomatoid fungi on various tree species in Norway and Poland, three unusual species, which fit the broader morphological description of Leptographium spp., were found in association with Scolytus ratzeburgi, Dryocoetes alni and Trypodendron domesticum on a variety of hardwoods, and from wounds on Tilia cordata. Phylogenetic analyses of sequence data for three gene regions (ITS2-LSU, β-tubulin, and TEF1-α) showed that these Leptographium species are phylogenetically closely related to each other and form a well-supported lineage that included Grosmannia grandifoliae and Leptographium pruni. The first species could be distinguished from the other Leptographium species based on conidiophores arising from spiral hyphae, chlamydospore-like structures and a hyalorhinocladiella-like synanamorph in culture. The second species differs from the previous one by having distinctly shorter conidiophores and smaller conidia. This species also produces a well-developed sporothrix-like synanamorph with denticulate conidiogenous cells. Based on these unusual morphological characteristics and distinct DNA sequences, these fungi were recognised as new taxa for which the names Leptographium trypodendri sp. nov. and L. betulae sp. nov. are provided. The third group of isolates belonged to Grosmannia grandifoliae, representing the first report of this species outside of the USA. The newly defined G. grandifoliae complex is the first species complex in Leptographium s.l. consisting of only hardwood-infecting species.
Authors
Leif Sundheim Åshild Ergon Christer Magnusson Jan Netland Egil Prestløkken Arild Sletten May Sæthre Elin Thingnæs Lid Tron Øystein Gifstad Micael Wendell Guro Brodal Halvor Solheim Anne Marte Tronsmo Bjørn Økland Trond RafossAbstract
No abstract has been registered
Authors
Ari Hietala Hugh Cross Halvor Solheim Volkmar Timmermann Nina Elisabeth Nagy Isabella Børja Adam Vivian-Smith Jørn Henrik SønstebøAbstract
No abstract has been registered
Authors
Hugh Cross Jørn Henrik Sønstebø Nina Elisabeth Nagy Volkmar Timmermann Halvor Solheim Isabella Børja Håvard Kauserud Tor Carlsen Barbara Rzepka Katarzyna Wasak Adam Vivian-Smith Ari HietalaAbstract
High biodiversity is regarded as a barrier against biological invasions. We hypothesized that the invasion success of the pathogenic ascomycete Hymenoscyphus fraxineus threatening common ash in Europe relates to differences in dispersal and colonization success between the invader and the diverse native competitors. Ash leaf mycobiome was monitored by high-throughput sequencing of the fungal internal transcribed spacer region (ITS) and quantitative PCR profiling of H. fraxineus DNA. Initiation of ascospore production by H. fraxineus after overwintering was followed by pathogen accumulation in asymptomatic leaves. The induction of necrotic leaf lesions coincided with escalation of H. fraxineus DNA levels and changes in proportion of biotrophs, followed by an increase of ubiquitous endophytes with pathogenic potential. H. fraxineus uses high propagule pressure to establish in leaves as quiescent thalli that switch to pathogenic mode once these thalli reach a certain threshold – the massive feedback from the saprophytic phase enables this fungus to challenge host defenses and the resident competitors in mid-season when their density in host tissues is still low. Despite the general correspondence between the ITS-1 and ITS-2 datasets, marker biases were observed, which suggests that multiple barcodes provide better overall representation of mycobiomes.
Abstract
The root rot pathogens in Norway spruce (Picea abies) Heterobasidion ssp. cause substantial loss in carbon sequestered in forest and economic revenue for forest owners. To facilitate strategic breeding planning for increased resistance against this pathogen in particular, the blue stain fungus Endoconidiophora polonica, growth and wood quality traits (wood density and spiral grain), we estimated additive genetic parameters, correlations and the potential response from selection. Parameters were estimated from a progeny trial series established at two sites (25 years from planting) and their parents in a seed orchard (43 years from grafting). A standard half-sib analysis based on progenies and a parent–offspring regression was used for estimation of heritabilities. Resistance against the pathogens was measured as lesion length under bark after inoculations in phloem. Heritability values varied with site and estimation procedure from 0.06 to 0.33, whereas the phenotypic variance (as CV P ) is high and fairly stable around 40–50 %. Heritability values for wood density and spiral grain in the same material varied from 0.32 to 0.63. The highest heritability values were generally obtained from parent–offspring regression. There is no evidence of resistance traits being genetically correlated with growth or wood quality traits. Wood density is negatively correlated with stem diameter. Implications for breeding are discussed.
Authors
Rein Drenkhan V Tomesova-Haataja S Fraser RE Bradshaw P Vahalik MS Mullett J Martin-Garcia LS Bulman MJ Wingfield T Kirisits TL Cech S Schmitz R Baden K Tubby A Brown M Georgieva A Woods R Ahumada L Jankovsky IM Thomsen K Adamson B Marcais M Vuorinen P Tsopelas A Koltay A Halasz N La Porta N Anselmi R Kiesnere S Markovskaja A Kacergius I Papazova-Anakieva M Risteski K Sotirovski J Lazarevic Halvor Solheim P Boron H Braganca D Chira DL Musolin AV Selikhovkin TS Bulgakov N Keca D Karadzic V Galovic P Pap M Markovic L. Poljakovic Pajnik V Vasic E Ondruskova B Piskur D Sadikovic JJ Diez A Solla H Millberg J Stenlid A Angst V Queloz A Lehtijärvi HT Dogmus-Lehtijärvi F Oskay K Davydenko V Meshkova D Craig S Woodward Irene BarnesAbstract
Dothistroma needle blight (DNB) is one of the most important diseases of pine. Although its notoriety stems from Southern Hemisphere epidemics in Pinus radiata plantations, the disease has increased in prevalence and severity in areas of the Northern Hemisphere, including Europe, during the last two decades. This increase has largely been attributed to expanded planting of susceptible hosts, anthropogenic dispersal of the causative pathogens and changes in climate conducive to disease development. The last comprehensive review of DNB was published in 2004, with updates on geographic distribution and host species in 2009. Importantly, the recognition that two species, Dothistroma septosporum and D. pini, cause DNB emerged only relatively recently in 2004. These two species are morphologically very similar, and DNA-based techniques are needed to distinguish between them. Consequently, many records of host species affected or geographic location of DNB prior to 2004 are inconclusive or even misleading. The objectives of this review were (i) to provide a new database in which detailed records of DNB from 62 countries are collated; (ii) to chart the current global distribution of D. septosporum and D. pini; (iii) to list all known host species and to consider their susceptibility globally; (iv) to collate the published results of provenance trials; and (v) to consider the effects of site factors on disease incidence and severity. The review shows that DNB occurs in 76 countries, with D. septosporum confirmed to occur in 44 and D. pini in 13. There are now 109 documented Pinaceae host taxa for Dothistroma species, spanning six genera (Abies, Cedrus, Larix, Picea, Pinus and Pseudotsuga), with Pinus being the dominant host genus, accounting for 95 host taxa. The relative susceptibilities of these hosts to Dothistroma species are reported, providing a resource to inform species choice in forest planting. Country records show that most DNB outbreaks in Europe occur on Pinus nigra and its subspecies. It is anticipated that the collaborative work described in this review will both underpin a broader global research strategy to manage DNB in the future and provide a model for the study of other forest pathogens.
Authors
R. Drenkhan Halvor Solheim A. Bogacheva T. Riit K. Adamson T. Drenkhan T. Maaten Ari HietalaAbstract
Dieback of European ash was first observed in Europe in the early 1990s. The disease is caused by the invasive ascomycete Hymenoscyphus fraxineus, proposed to originate from Far East Asia, where it has been considered a harmless saprotroph. This study investigates the occurrence of H. fraxineus in tissues of local ash species in the Russian Far East, and assesses its population-specific genetic variation by ITS sequencing. Shoot dieback symptoms, characteristic of H. fraxineus infection on European ash, were common, but not abundant, on Fraxinus mandshurica and Fraxinus rhynchophylla trees in Far East Russia. High levels of pathogen DNA were associated with necrotic leaf tissues of these ash species, indicating that the local H. fraxineus population is pathogenic to their leaves. However, the low levels of H. fraxineus DNA detected in shoots with symptoms, the failure to isolate this fungus from such tissues, and the presence of other fungi with pathogenic potential in shoots with symptoms indicate that local H. fraxineus strains may not be responsible (or their role is negligible) for the observed ash shoot dieback symptoms in the region. Conspicuous differences in ITS rDNA sequences detected between H. fraxineus isolates from Russian Far East and European populations suggest that the current ash dieback epidemic in Europe might not directly originate from the Russian Far East. Revision of the herbarium material shows that the earliest specimen of H. fraxineus was collected in 1962 from the Russian Far East and the oldest H. fraxineus specimen of China was collected in 2004.
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Authors
Riikka Linnakoski Robert Jankowiak Caterina Villari Thomas Kirisits Halvor Solheim Z. Wilhelm de Beer Michael J. WingfieldAbstract
Two species of blue-stain fungi with similar morphologies, Ophiostoma brunneo-ciliatum and Ophiostoma clavatum, are associates of bark beetles infesting Pinus spp. in Europe. This has raised questions whether they represent distinct taxa. Absence of herbarium specimens and contaminated or mistakenly identified cultures of O. brunneo-ciliatum and O. clavatum have accentuated the uncertainty regarding their correct identification. The aim of this study was to reconsider the identity of European isolates reported as O. brunneo-ciliatum and O. clavatum by applying DNA-based identification methods, and to provide appropriate type specimens for them. Phylogenetic analyses of the ITS, βT, TEF-1α and CAL gene sequences revealed that the investigated isolates represent a complex of seven cryptic species. The study confirmed that ITS data is insufficient to delineate species in some Ophiostoma species clusters. Lectotypes and epitypes were designated for O. clavatum and O. brunneo-ciliatum, and three new species, Ophiostoma brunneolum, Ophiostoma macroclavatum and Ophiostoma pseudocatenulatum, are described in the newly defined O. clavatum-complex. The other two species included in the complex are Ophiostoma ainoae and Ophiostoma tapionis. The results suggest co-evolution of these fungi in association with specific bark beetles. The results also confirm the identity of the fungus associated with the pine bark beetle Ips acuminatus as O. clavatum, while O. brunneo-ciliatum appears to be mainly associated with another pine bark beetle, Ips sexdentatus.
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Globally, billions of tons of carbon sequestered in trees are annually recycled back to the atmosphere through wood decomposition by microbes. In Norway, every fifth Norway spruce shows at final harvest infection by pathogenic white-rot fungi in the genera Heterobasidion and Armillaria. As these fungi can mineralize all components of wood, we predicted that they have a significant carbon footprint. Gas samples taken from infected stems were analyzed for CO2 and CH4 concentrations, and wood samples from different parts of the decay columns were incubated under hypoxic (4% O2) and anoxic laboratory conditions. In spring and summer the stem concentrations of CO2 were generally two times higher in trees with heartwood decay than in healthy trees. For most of the healthy trees and trees with heartwood decay, mean stem concentrations of CH4 were comparable to ambient air, and only some Armillaria infected trees showed moderately elevated CH4. Consistently, low CH4 production potentials were recorded in the laboratory experiment. Up-scaling of CO2 efflux due to wood decay in living trees suggests that the balance between carbon sequestration and emission may be substantially influenced in stands with high frequency of advanced root and stem heartwood decay.
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Halvor SolheimAbstract
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Ari Hietala Isabella Børja Hugh Cross Volkmar Timmermann Nina Elisabeth Nagy Halvor Solheim Jørn Henrik Sønstebø adam vivian smithAbstract
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Ari Hietala Isabella Børja Hugh Cross Nina Elisabeth Nagy Halvor Solheim Jørn Henrik Sønstebø Volkmar Timmermann Adam Vivian-SmithAbstract
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Ari Hietala Hugh Cross Isabella Børja Nina Elisabeth Nagy Volkmar Timmermann Halvor Solheim Jørn Henrik Sønstebø Adam Vivian-SmithAbstract
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Ten saplings of European ash (Fraxinus excelsior L.) naturally infected by the invasive ash dieback pathogen Hymenoscyphus fraxineus were collected in Ukraine and Norway and examined for bark necrosis and extension of discoloration in sapwood and pith in a stem region. Tissue-specific colonization profiles were determined by spatial analyses of symptomatic and visually healthy stem tissues using a H. fraxineus-specific qPCR assay and light microscopy. Our data suggest that hyphal growth in the starch-rich perimedullary pith is of particular importance for both axial and radial spread of H. fraxineus, but that most of its biomass accumulates in sapwood parenchyma. The study confirms the results from earlier work and presents new information that refines the current stem invasion model.
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Jonas Oliva S. Rommel Carl Gunnar Fossdal Ari Hietala M. Nemesio-Gorriz Halvor Solheim Malin ElfstrandAbstract
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Halvor SolheimAbstract
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Leif Sundheim Trond Hofsvang Christer Magnusson Gunnar Sundstøl Eriksen Lars Olav Brandsæter Guro Brodal Siv Fagertun Remberg Anne Kjersti Uhlen Åshild Kristine Andreassen Augustine Arukwe Aksel Bernhoft Knut Egil Bøe Margaretha Haugen Gro Ingunn Hemre Åshild Krogdahl Torsten Källqvist Jørgen Fredrik Lassen Bjørn Næss Trond Rafoss Janneche Utne Skåre Arild Sletten Halvor Solheim Inger-Lise Steffensen Ole Torrissen Anne Marte Tronsmo Bjørn Økland Jan AlexanderAbstract
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Ari M. Hietala, Volkmar Timmermann, Isabella Børja & Halvor Solheim Norwegian Forest and Landscape Institute. PO Box 115, 1431 Ås, Norway: ari.hietala@skogoglandskap.no Owing to the Gulf Stream, the northernmost European populations of several tree species are found in Norway. Common ash (Fraxinus excelsior), the only native ash species in Norway, is present in the lowlands in the southeastern part with continental climate and in southern and southwestern coastal regions with North Atlantic climate up to Central Norway. The current standing volume of ash in Norway is ca 3 mill m3 (broadleaved trees in total 220 mill m3). The first documentation of Ash Dieback (ADB) is from 2008 from a nursery in the southeastern part of the country. A survey later that year showed that dieback symptoms were present over a distance of nearly 400 km in the southeastern region. In addition to nurseries and forests, ADB symptoms were observed on roadside, alley, garden and park trees. Based on the presence of old ADB-like stem lesions detected in 2008, the pathogen must have arrived to Norway no later than 2006. In 2008, the Norwegian Food Safety Authority laid down regulations with the aim of preventing further spread of ADB. These regulations divide the country into quarantine, observation and infection-free zones, and prohibit the export of ash seedlings, seed and wood from the quarantine zone. Despite of these regulations, the disease spread rapidly along the western coast in the period between 2009 and 2013, and currently only the ash stands in Central Norway are free of the disease. The rapid spread of the disease in Norway is obviously due to airborne dispersal of pathogen ascospores. In our experimental stand in SE Norway the number of pathogen fruit bodies can be as high as 10,000 per m2 in the peak season, mid-July to mid-August. During the early morning hours the amount of pathogen ascospores at a diseased stand can exceed 100,000 ascospores per m3 air. The first symptoms of the disease, necrotic lesions on leaf blade and petiole, appear typically during the first two weeks of August in SE Norway. To observe long-term impacts of ADB, eight monitoring plots have been established in continental and North Atlantic climate zones. In SE Norway with the oldest disease history, above 60 % of the trees with a breast height diameter (BHD) below 12.5 cm have so far died or suffer from severe defoliation, 1/3 of the larger trees being affected to a similar degree. The proportions of healthy (no signs of defoliation) small and larger trees are 20% and 37%, respectively. In SW Norway with more recent disease history a similar trend is observed but the proportion of dead trees is still small. As a consequence of ADB, the Norwegian nurseries no longer grow ash seedlings. There are currently no practical control options for the disease in forestland. Several European countries have reported that even at heavily diseased ash stands there are often some ash trees that show little symptoms. This may be due to genetic variation between trees in disease resistance, a hypothesis that is currently being investigated in several European projects. Thus implementation of forest management practices that eliminate ash could have a negative effect as survival of the tree ultimately depends on selection of trees with increased disease resistance. Bibliography for Ari M. Hietala Ari M. Hietala is a Senior Forest Pathologist at the Norwegian Forest and Landscape Institute, which is a primarily government funded organisation providing scientific research and services to government, non-governmental and commercial organisations. He has worked with a range of fungal root and shoot diseases occurring on broadleaved trees and conifers indigenous to the Nordic countries. Ari and the rest of the group participate currently in several European consortia engaged in ash dieback research.
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Daniel Flø Halvor Solheim Christer Magnusson Trond Rafoss Leif Sundheim Juliana Irina Spies PerminowAbstract
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Leif Sundheim Daniel Flø Trond Rafoss Guro Brodal Åshild Ergon Christer Magnusson Arild Sletten Halvor Solheim May Sæthre Anne Marte Tronsmo Bjørn ØklandAbstract
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Nina Elisabeth Nagy Katarzyna Sikora Paal Krokene Ari Hietala Halvor Solheim Carl Gunnar FossdalAbstract
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Igor A. Yakovlev Ari Hietala Pierre-Emmanuel Courty Taina Lundell Halvor Solheim Carl Gunnar FossdalAbstract
The pathogenic white-rot basidiomycete Heterobasidion irregulare is able to remove lignin and hemicellulose prior to cellulose during the colonization of root and stem xylem of conifer and broadleaf trees. We identified and followed the regulation of expression of genes belonging to families encoding ligninolytic enzymes. In comparison with typical white-rot fungi, the H. irregulare genome has exclusively the short-manganese peroxidase type encoding genes (6 short-MnPs) and thereby a slight contraction in the pool of class II heme-containing peroxidases, but an expansion of the MCO laccases with 17 gene models. Furthermore, the genome shows a versatile set of other oxidoreductase genes putatively involved in lignin oxidation and conversion, including 5 glyoxal oxidases, 19 quinone-oxidoreductases and 12 aryl-alcohol oxidases. Their genetic multiplicity and gene-specific regulation patterns on cultures based on defined lignin, cellulose or Norway spruce lignocellulose substrates suggest divergent specificities and physiological roles for these enzymes. While the short-MnP encoding genes showed similar transcript levels upon fungal growth on heartwood and reaction zone (RZ), a xylem defense tissue rich in phenolic compounds unique to trees, a subset of laccases showed higher gene expression in the RZ cultures. In contrast, other oxidoreductases depending on initial MnP activity showed generally lower transcript levels on RZ than on heartwood. These data suggest that the rate of fungal oxidative conversion of xylem lignin differs between spruce RZ and heartwood. It is conceivable that in RZ part of the oxidoreductase activities of laccases are related to the detoxification of phenolic compounds involved in host-defense. Expression of the several short-MnP enzymes indicated an important role for these enzymes in effective delignification of wood by H. irregulare.
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Leif Sundheim Christer Magnusson Trond Rafoss Halvor Solheim Bjørn Økland Trond Hofsvang Arild Sletten Anne Marte Tronsmo Daniel FløAbstract
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Authors
Halvor SolheimAbstract
Common juniper (Juniperus communis) hosts not many pests or pathogens, but recently increasing needle blight has been observed in Norway. During a survey the needle blight was recorded in many parts of southern Norway but not above 550 m a.s.l., and it has been found both in forests, pastures and gardens. Trees are affected differently; some trees seem to be unaffected, while other trees may be killed. The cause of the disease is a fungus in the family Mycosphaerellaceae hitherto not reported from Norway. In forest pathology literature it has been named Stigmina juniperina, but also Asperisporium juniperinum. However, based on results of molecular sequence analyses it is proposed here that a more appropriate name should be Passalora juniperina (Georgescu & Badea) H. Solheim comb. Nov.
Authors
Halvor SolheimAbstract
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Authors
Aksel Bernhoft Gunnar Sundstøl Eriksen Leif Sundheim Marc Berntssen Anne Lise Brantsæter Guro Brodal Christiane Kruse Fæste Ingerd Skow Hofgaard Trond Rafoss Tore Sivertsen Anne Marte Tronsmo Heidi Amlund Augustine Arukwe Marit Aursand Margaretha Haugen Gro Ingunn Hemre Trond Hofsvang Helle Katrine Knutsen Åshild Krogdahl Jørgen Fredrik Lassen Christer Magnusson Audun Helge Nerland Live Lingaas Nesse Bjørn Næss Einar Ringø Anders Ruus Janneche Utne Skåre Halvor Solheim Arild Sletten Inger-Lise Steffensen Line Charlotte Sverdrup Birger Svihus Ole Torrissen Bente Elisabeth Torstensen Cathrine Thomsen Robin Ørnsrud Bjørn Økland Olav Østerås Jan AlexanderAbstract
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A survey to identify ophiostomatoid fungi that infect wounds on native Norwegian and Swedish broadleaved trees was undertaken during summer 2004. A fungus resembling a species of Sporendocladia was commonly isolated from the exposed cambium and inner bark of wounds. Morphological examination and comparisons of DNA sequence data for the ITS and 5.8S regions of the rRNA gene region led to its identification as Sporendocladia bactrospora. Pathogenicity trials on young Populus tremula and Betula pubescens trees showed that S. bactrospora is capable of causing lesions on these trees. There have been few previous reports of S. bactrospora, and in most cases, these have been as saprophytes on wood. In contrast, results of this study show that it is a common inhabitant of freshly made wounds on native broadleaved trees in Scandinavia, and it appears to contribute to staining of wood.
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Shoot dieback disease of European ash caused by the ascomycete Hymenoscyphus pseudoalbidus threatens ash on a continental scale. A spore sampler placed in a diseased ash forest in Southern Norway, coupled with microscopy and DNA-based fungal species-specific real-time PCR assays, was employed to profile diurnal and within-season variation in infection pressure by ascospores of H. pseudoalbidus and the potentially co-existing non-pathogenic Hymenoscyphusalbidus. Hymenoscyphus pseudoalbidus was found to be predominant in the stand. Massive simultaneous liberation, by active discharge of pathogen ascospores in the morning, peaked in mid-Jul. to mid-Aug. Accumulation of pathogen DNA on leaflets of current-year leaves reached a high level plateau phase before appearance of autumn coloration, suggesting that pathogen establishment in leaves is terminated before the onset of leaf senescence.
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Winter damage caused by frost is frequently observed on common ash (Fraxinus excelsior) in Norway. In spring 2007, extensive winter damage most likely camouflaged ash dieback caused by Chalara fraxinea. In 2008, ash dieback caused by C. fraxinea had spread to large areas in the southern part of Norway. The disease was widespread in forests and nurseries, but also on roadside trees, and in gardens and parks. In 2009, the disease had spread to new areas; about 30 km into Rogaland county in southwestern Norway and also further into some valleys in southeastern Norway.
Authors
Alberto Santini Luisa Ghelardini Ciro De Pace Marie-Laure Desprez-Loustau Paolo Capretti Anne Chandelier Thomas Cech Danut Chira Stephanos Diamandis Talis Gaitnieks Jarkko Hantula Ottmar Holdenrieder Libor Jankovský Thomas Jung Dusan Jurc Thomas Kirisits Andrej Kunca Vaidotas Lygis Monika Malecka Benoit Marcais Sophie Schmitz Jörg Schumacher Halvor Solheim Alejandro Solla Ilona Szabò Panaghiotis Tsopelas Andrea Vannini Anna Maria Vettraino Joan Webber Stephen Woodward Jan StenlidAbstract
A large database of invasive forest pathogens (IFPs) was developed to investigate the patterns and determinants of invasion in Europe. Detailed taxonomic and biological information on the invasive species was combined with country-specific data on land use, climate, and the time since invasion to identify the determinants of invasiveness, and to differentiate the class of environments which share territorial and climate features associated with a susceptibility to invasion. IFPs increased exponentially in the last four decades. Until 1919, IFPs already present moved across Europe. Then, new IFPs were introduced mainly from North America, and recently from Asia. Hybrid pathogens also appeared. Countries with a wider range of environments, higher human impact or international trade hosted more IFPs. Rainfall influenced the diffusion rates. Environmental conditions of the new and original ranges and systematic and ecological attributes affected invasiveness. Further spread of established IFPs is expected in countries that have experienced commercial isolation in the recent past. Densely populated countries with high environmental diversity may be the weakest links in attempts to prevent new arrivals. Tight coordination of actions against new arrivals is needed. Eradication seems impossible, and prevention seems the only reliable measure, although this will be difficult in the face of global mobility.
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The relative frequency of Therrya fuckelii and T. pini fruiting on dead branches of Scots pine was investigated in southern Norway by examining lightning-damaged and wind-fallen trees, randomly collected branches and Nordic herbarium collections of these ascomycetes representing the order Rhytismatales. Ascus, ascospore, and subhymenium characteristics were used as criteria for species identification, while a sequence analysis of ITS rDNA gene cluster was performed to compare the relatedness of the species to each other and to corresponding fungal sequences available at the NCBI GenBank Sequence Database. In a few cases, the two Therrya species co-occurred on the same branch, but in general, whether field or herbarium material, T. fuckelii was clearly more common than T. pini.Within the Nordic countries, both species occurred throughout the natural distribution area of Scots pine. The ITS rDNA sequence of T. pini strains was 91% similar to T. fuckelii strains, the differences locating both within the internal transcribed spacers ITS1 and ITS2 and the 5.8 S rDNA gene. More variation in the ITS1-5.8S-ITS2 sequence was observed among T. pini than T. fuckelii samples; genetic implications of this finding are discussed. Upon sequence analysis, we discovered that a T. pini sequence has been deposited in the NCBI GenBank under a false identity. We emphasize the importance of co-examining strains that originate from mature fruit bodies with fully developed morphologic features as reference samples.
Authors
Nadeem Yaqoob Igor A. Yakovlev Paal Krokene Harald Kvaalen Halvor Solheim Carl Gunnar FossdalAbstract
We compared gene expression in Norway spruce secondary phloem (bark) and developing xylem (sapwood) in response to the necrotrophic pathogen Heterobasidion parviporum, wounding and methyl jasmonate (MeJ). The pathogen induced systemic and local up-regulation of PaPX3, PaPX2 and PaChi4 in both bark and sapwood that returned to constitutive levels as the plants recovered from the infection, whereas the local responses to MeJ were similar in both tissues but was longer lasting for PaPX3 and PaChi4. Genes involved in lignin biosynthesis (PaPAL1, PaPAL2, PaC4H3/5 and PaHCT1) were up-regulated locally in the bark in response to pathogen and wounding whereas MeJ induced a similar but stronger local response. The ethylene biosynthesis related transcripts PaACO and PaACS did not increase in response to MeJ treatment or the pathogen, however it increased both locally and systemically as a response to wounding in the sapwood. These results demonstrate that the local and systemic host responses to pathogen infection and wounding largely correspond and reveal striking similarities between the local response to a necrotroph, wounding and MeJ treatment in both bark and living wood.
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Ceratocystis polonica and Heterobasidion parviporum are important fungal pathogens in Norway spruce (Picea abies). Tree susceptibility to these pathogens with respect to phenology was studied using artificial fungal inoculations at six stages of bud development, and assessed by measuring phloem necroses in the stems of 2- and 8-year-old trees. Tree capacity for resistance was assessed by measuring phloem nonstructural carbohydrates at each stage. Phloem necroses were significantly larger in trees with fungal versus control inoculations and increased significantly over time. Changes in nonstructural carbohydrates occurred in the trees; a significant decline in starch and a slight but significant increase in total sugars occurred over time. These results suggest that susceptibility to fungal pathogens and carbohydrate levels in the stems of the trees were related to fine-scale changes in bud development. A trade-off may occur between allocation of starch (the major fraction of the stem carbohydrate pool) to bud development/shoot growth versus defence of the stem. Previous tests of plant defence hypotheses have focused on herbivory on plants growing under different environmental conditions, but the role of phenology and the effect of pathogens are also important to consider in understanding plant resource allocation patterns.
Authors
Tao Zhao Paal Krokene Niklas Björklund Nadir Erbilgin Erik Christiansen Bo Långström Halvor Solheim Anna-Karin Borg-KarlsonAbstract
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Carl Gunnar Fossdal Nina Elisabeth Nagy Ari Hietala Harald Kvaalen Rune Slimestad Steve Woodward Halvor SolheimAbstract
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Authors
Åke Olson Andrea Aerts Fred Asiegbu Lassaad Belbahri Ourdia Bouzid Anders Broberg Björn Canback Pedro M. Coutinho Dan Cullen Kerstin Dalman Giuliana Deflorio Linda T.A. van Diepen Christophe Dunand Sébastien Duplessis Mikael Durling Paolo Gonthier Jane Grimwood Carl Gunnar Fossdal David Hansson Bernard Henrissat Ari Hietala Kajsa Himmelstrand Dirk Hoffmeister Nils Högberg Timothy Y. James Magnus Karlsson Annegret Kohler Ursula Kües Yong-Hwan Lee Yao-Cheng Lin Mårten Lind Erika Lindquist Vincent Lombard Susan Lucas Karl Lundén Emmanuelle Morin Claude Murat Jongsun Park Tommaso Raffaello Pierre Rouzé Asaf Salamov Jeremy Schmutz Halvor Solheim Jerry Ståhlberg Heriberto Vélëz Ronald P. de Vries Ad Wiebenga Steve Woodward Igor A. Yakovlev Matteo Garbelotto Francis Martin Igor V. Grigoriev Jan StenlidAbstract
• Parasitism and saprotrophic wood decay are two fungal strategies fundamental for succession and nutrient cycling in forest ecosystems. An opportunity to assess the trade-off between these strategies is provided by the forest pathogen and wood decayer Heterobasidion annosum sensu lato. • We report the annotated genome sequence and transcript profiling, as well as the quantitative trait loci mapping, of one member of the species complex: H. irregulare. Quantitative trait loci critical for pathogenicity, and rich in transposable elements, orphan and secreted genes, were identified. • A wide range of cellulose-degrading enzymes are expressed during wood decay. By contrast, pathogenic interaction between H. irregulare and pine engages fewer carbohydrate-active enzymes, but involves an increase in pectinolytic enzymes, transcription modules for oxidative stress and secondary metabolite production. • Our results show a trade-off in terms of constrained carbohydrate decomposition and membrane transport capacity during interaction with living hosts. Our findings establish that saprotrophic wood decay and necrotrophic parasitism involve two distinct, yet overlapping, processes.
Authors
Carl Gunnar Fossdal Nadeem Yaqoob Paal Krokene Harald Kvaalen Halvor Solheim Igor A. YakovlevAbstract
Background: NB-LRR resistance proteins are involved in recognizing pathogens and other exogenous stressors in plants. Resistance proteins are the first step in induced defence responses and a better understanding of their regulation is important to understand the mechanisms of plant defence. Much of the post-transcriptional regulation in plants is controlled by microRNAs (miRNA). We examined the expression of five Norway spruce miRNA that may regulate NB-LRR related transcripts in secondary phloem (bark) of resistant Norway spruce after wounding and inoculation with the necrotrophic blue stain fungus Ceratocystis polonica. Results: The plants of this clone recovered from both the pathogen inoculations and wounding alone. We found local and systemic induction of the resistance marker genes PaChi4, PaPAL and PaPX3 indicative of an effective induced host defence response. There were minor local and systemic changes in the expression of five miRNAs and 21 NB-LRRs between healthy and treated plants. Only five putative NB-LRRs (PaLRR1, PaLRR3, PaLRR14, PaLRR15 and PaLRR16) showed significant increases greater than two-fold as a local response to C. polonica. Of all NB-LRRs only PaLRR3, the most highly differentially regulated NB-LRR, showed a significant increase also due to wounding. The five miRNAs showed indications of an initial local and systemic down-regulation at day 1, followed by a later increase up to and beyond the constitutive levels at day 6. However, the initial down-regulation was significant only for miR3693 and miR3705. Conclusions: Overall, local and systemic expression changes were evident only for the established resistance marker genes and PaLRR3. The minor expression changes observed both for the followed miRNAs and their predicted NB-LRR targets suggest that the expression of most NB-LRR genes are maintained close to their constitutive levels in stressed and healthy Norway spruce plants.
Authors
Arild Sletten Trond Hofsvang Christer Magnusson Trond Rafoss Halvor Solheim Anne Marte Tronsmo Bjørn Økland Leif SundheimAbstract
No abstract has been registered
Abstract
The ascomycete fungus Hymenoscyphus pseudoalbidus (anamorph Chalara fraxinea) is responsible for ash dieback currently expanding over large parts of Europe. Our objective was to investigate the genetic structure of H. pseudoalbidus and to examine its relationship to the species H. albidus, known as a saprotroph. The study comprised 181 isolates of H. pseudoalbidus collected within the diseased area, 17 H. albidus isolates from six apothecia, collected outside the diseased area in Norway, and nine apothecia of H. pseudoalbidus collected in Sweden. By analysis of microsatellite markers developed for this study, combined with AP-PCR using the M13 primer, we demonstrated sexual heterothally in H. pseudoalbidus, detected high gene flow and low geographic structure of the H. pseudoalbidus population and found indications of a founder effect. Also, substantial genetic differences were detected between the two species of fungi; only four of seven microsatellite markers developed for H. pseudoalbidus were amplified for H. albidus, and no alleles were shared among the species. Furthermore, AP-PCR banding patterns were distinctly different for the two species. We conclude that even though the two fungi have a similar habitat and are morphologically virtually identical, they do not share a recent common ancestor.
Abstract
Aspen trees are exposed to a range of attackers and employ varied strategies to reduce their impact. The diversity of responses may have importance for resistance properties at the stand level, and justifies the search for varied defensive strategies in natural populations. We used transcriptomic tools to evaluate diverse responses at the gene activity level in Populus tremula in response to wounding, and to inoculation with two pathogenic fungi (Melampsora magnusiana vs Ceratocystis sp.) that differ in life style (biotroph vs necrotroph) and host tissue requirement (live leaf vs dead wood tissues). Two aspen genotypes from the SwAsp collection with differences in growth and phenolic composition were used to study differences in resistance properties. High defence gene induction, high growth and elevated defence properties toward the biotroph appeared to support each other in this study exemplified in the more resistant SwAsp clone, whereas the more susceptible SwAsp clone was much less responsive to infections, and displayed more symptoms when infected with M. magnusiana. Interestingly, in the more resistant clone wounding gave greater systemic activity of selected candidate genes than when combined with the necrotroph, suggesting that this pathogen has some ability to suppress the induction or translocation of the systemic defence signal in this particular clone.
Authors
Carl Gunnar Fossdal Ari M. Hietala Igor A. Yakovlev Gustav Vaaje-Kolstad Emil Stefaczyk Halvor SolheimAbstract
The GH61 represents the most enigmatic Glycoside Hydrolase Family (GH) regarding putative enzymatic activity and importance in cellulose degradation. Heterobasidion irregulare is a necrotizing pathogen and white rot fungus, causing enormous damages in conifer forests.The genome of H. irregulare allowed identification of ten HiGH61 genes. qRT-PCR analysis separate the HiGH61 members into two groups; one that show up regulation on lignocellulosic substrates and another that show either down regulation or constitutive expression. This grouping suggests that the fungus relates different sets of GH61s for different substrates, like in the various stages of necrotizing and saprophytic growth on the host.One HiGH61 showed up to 17000 fold increase on spruce heartwood suggesting a pivotal role in cellulose decomposition during saprophytic growth. Sequence analysis of these genes reveals that all GH61s but one possess the conserved metal binding motif predicted to be essential for activity.The sequences also divide into groups having either an insert near the N-terminus or an insert near the second catalytic histidine, which both may represent extensions of the substrate binding surface. Three HiGH61s encode cellulose-binding modules (CBM1), indicating direct targeting of crystalline cellulose, two being up regulated on pure cellulose.There was a common substrate-specific induction patterns of the HiGH61s with several reference cellulolytic and hemicellulolytic GHs, this taken together with their low levels on media lacking lignocellulose, reflect the concerted nature of cell wall polymer degradation.
Authors
Igor A. Yakovlev Gustav Vaaje-Kolstad Ari Hietala Emil Stefanczyk Halvor Solheim Carl Gunnar FossdalAbstract
No abstract has been registered
Authors
Leif Sundheim Daniel Flø Trond Rafoss Trond Hofsvang Christer Magnusson Arild Sletten Halvor Solheim Anne Marte Tronsmo Bjørn ØklandAbstract
No abstract has been registered
Authors
Hugo Germain Denis Lachance Gervais Pelletier Carl Gunnar Fossdal Halvor Solheim Armand SeguinAbstract
A 1149 bp genomic fragment corresponding to the 5' non-coding region of the PgD1 (Picea glauca Defensin 1) gene was cloned, characterized, and compared with all Arabidopsis thaliana defensin promoters. The cloned fragment was found to contain several motifs specific to defence or hormonal response, including a motif involved in the methyl jasmonate reponse, a fungal elicitor responsive element, and TC-rich repeat cis-acting element involved in defence and stress responsiveness. A functional analysis of the PgD1 promoter was performed using the uidA (GUS) reporter system in stably transformed Arabidopsis and white spruce plants. The PgD1 promoter was responsive to jasmonic acid (JA), to infection by fungus and to wounding. In transgenic spruce embryos, GUS staining was clearly restricted to the shoot apical meristem. In Arabidopsis, faint GUS coloration was observed in leaves and flowers and a strong blue colour was observed in guard cells and trichomes. Transgenic Arabidopsis plants expressing the PgD1::GUS construct were also infiltrated with the hemibiotrophic pathogen Pseudomonas syringae pv. tomato DC3000. It caused a suppression of defensin expression probably resulting from the antagonistic relationship between the pathogen-stimulated salicylic acid pathway and the jasmonic acid pathway. It is therefore concluded that the PgD1 promoter fragment cloned appears to contain most if not all the elements for proper PgD1 expression and that these elements are also recognized in Arabidopsis despite the phylogenetic and evolutionary differences that separates them.
Authors
Nina Elisabeth Nagy Harald Kvaalen Monica Fongen Carl Gunnar Fossdal Nicholas Clarke Halvor Solheim Ari HietalaAbstract
Pathogen challenge of tree sapwood induces the formation of reaction zones with antimicrobial properties such as elevated pH and cation content. Many fungi lower substrate pH by secreting oxalic acid, its conjugate base oxalate being a reductant as well as a chelating agent for cations. To examine the role of oxalic acid in pathogenicity of white-rot fungi, we conducted spatial quantification of oxalate, transcript levels of related fungal genes, and element concentrations in heartwood of Norway spruce challenged naturally by Heterobasidion parviporum. In the pathogen-compromised reaction zone, upregulation of an oxaloacetase gene generating oxalic acid coincided with oxalate and cation accumulation and presence of calcium oxalate crystals. The colonized inner heartwood showed trace amounts of oxalate. Moreover, fungal exposure to the reaction zone under laboratory conditions induced oxaloacetase and oxalate accumulation, whereas heartwood induced a decarboxylase gene involved in degradation of oxalate. The excess level of cations in defense xylem inactivates pathogen-secreted oxalate through precipitation and, presumably, only after cation neutralization can oxalic acid participate in lignocellulose degradation. This necessitates enhanced production of oxalic acid by H. parviporum. This study is the first to determine the true influence of white-rot fungi on oxalate crystal formation in tree xylem.
Abstract
No abstract has been registered
Authors
Nina Elisabeth Nagy Simon Ballance Harald Kvaalen Carl Gunnar Fossdal Halvor Solheim Ari HietalaAbstract
Heterobasidion parviporum, a common pathogenic white-rot fungus in managed Norway spruce forests in northern and central Europe, causes extensive decay columns within stem heartwood of the host tree. Infected trees combat the lateral spread of decay by bordering the heartwood with a fungistatic reaction zone characterized by elevated pH and phenol content. To examine the mode of fungal feeding in the reaction zone of mature Norway spruce trees naturally infected by H. parviporum, we conducted spatial proWling of pectin and hemicellulose composition, and established transcript levels of candidate fungal genes encoding enzymes involved in degradation of the diVerent cell wall components of wood. Colonized inner heartwood showed pectin and hemicellulose concentrations similar to those of healthy heartwood, whereas the carbohydrate proWles of compromised reaction zone, irrespective of the age of fungal activity in the tissue, indicated selective fungal utilization of galacturonic acid, arabinose, xylose and mannose. These data show that the rate of wood decay in the reaction zone is slow. While the up-regulation of genes encoding pectinases and hemicellulases preceded that of the endoglucanase gene during an early phase of fungal interaction with xylem defense, the manganese peroxidase gene showed similar transcript levels during diVerent phases of wood colonization. It seems plausible that the reaction zone components of Norway spruce interfere with both lignin degradation and the associated co-hydrolysis of hemicelluloses and pectin, resulting in a prolonged phase of selective decay.
Abstract
Dieback of European ash (Fraxinus excelsior), caused by the ascomycete Hymenoscyphus pseudoalbidus (anamorph Chalara fraxinea), started around 1992 in Poland and has since then spread over large geographical areas. By November 2010, the disease had been recorded in 22 European countries. The gradual expansion and high intensity of the ash dieback epidemic in Europe may suggest that H. pseudoalbidus is an invasive alien organism. In Norway, ash dieback was first reported in spring 2008, and a survey in early summer of the same year revealed that the disease had spread over large parts of the southern and eastern regions of the country. The distance from the southernmost to the northernmost infected stands was, at that time, about 400 km. Some old necrotic lesions were also observed, indicating that the ash dieback pathogen is likely to have been present in Norway since at least 2006. In 2009, a spore sampler was installed in a diseased ash stand at Ås, South-Eastern Norway. Sampling started in late July and continued until late September. Large numbers of ascospores resembling those of H. pseudoalbidus were observed, with the maximum number of spores occurring from the end of July to mid-August. The deposition of ascospores occurred mainly between 6 and 8 a.m. Ascospores are most likely to be the primary source initiating host infections and responsible for the rapid recent spread of H. pseudoalbidus in Europe.
Abstract
A rapid increase in the frequency of Dutch elm disease (DED), a wilting disease of elm trees caused by bark-beetle vectored fungi, was observed in the early 1990s on several wych elm stands around Oslofjord, southern Norway. To examine the current status of the disease and its impacts on elm population, disease frequency and size distribution of elms were recorded at four locations. Northern parts of Lier, a municipality most affected by DED in Norway 15 years ago, showed in the survey season 4% disease frequency, whereas 13.8% of trees were dead, the dead trees having accumulated over several years in the unmanaged stands. In southern parts of the municipality the mean disease and death percentages were 1.9 and 2.4%. Compatible with their low disease incidence in early 1990s, the other two areas now examined, municipality of Larvik and district of Grenland, showed comparably low frequency of DED. Northern part of Lier showed significantly higher overall density of elm trees per hectare than the other examined areas, and also the small elms below 5 cm in d.b.h. were most frequent in this region. In contrast, the density of large trees was lower in northern Lier than in the other examined areas. These data suggest that regeneration of the tree is not prohibited owing to the disease but that the large trees have been locally reduced in frequency as a result of DED. The superior general density of elm trees in northern Lier, owing to the exceptionally rich soil in the warm southern slopes of the region,> may have contributed to the rapid increase of DED in the area 15 years ago and to the subsequent settlement of the disease outbreak as a chronic stage.
Abstract
The occurence of Armillaria species was assessed in Norway, enabling the northern-most distribution of this genus to be determined in Europe. Four Armillaria species were found in Norway. Armillaria borealis was the most common species occurring on woody vegetation to the permafrost zone (ca. 69°N). Armillaria cepistipes was present in southern and central Norway, but was not found further than 66°N. Armillaria solidipes and Armillaria gallica were rare, found at only one locality each; 59°40′ and 59°32′, respectively. Armillaria species were found on 14 hosts, but there was no significant difference between occurrence of A. borealis and A. cepistipes on declining and dead trees. Phylogenetic analyses separated each species into separate clades. All isolates of A. borealis, except one, and most isolates of A. solidipes were in separate clades. However, a subclade within the A. borealis clade was formed of two A. ostoyae and one A. borealis isolates. Two small A. cepistipes genets were found in a declining oak stand.
Abstract
During a survey conducted in August 2009 in northern Norway, symptoms typical for red band needle blight (1) were observed in four young Scots pine (Pinus sylvestris) stands. The stands, less than 15 years old, were located in humid sites near rivers in Bardu and Målselv municipalities, Troms County. Many of the oldest needles (2- to 3- years-old) in the lower part of young trees were partially or completely brown, but still attached, and red bands could be observed. Aggregations of conidial stromata were often seen in the red bands. Conidia were hyaline, smooth, thin walled, and filiform, 1.9 to 2.6 μm wide and 12 to 36 μm long....
Authors
Igor A. Yakovlev Ari M. Hietala Pierre-Emmanuel Courty Taina Lundell Jan Stenlid Halvor Solheim Carl Gunnar FossdalAbstract
In forest soils, saprotrophic, necrotrophic and ectomycorrhizal fungi are involved in carbon cycling. Heterobasidion annosum, white rot necrotrophic fungi, is known to decompose wood lignocellulose by secreting a broad range of oxidative enzymes. The genome H. annosum s.l. was sequenced by JGI to a 8.23X coverage and assembled into 39 scaffolds with a total size of 33.7 Mb covering more than 98% of the whole genome. Based on the genome sequence we have characterized gene families coding for enzymes known to participate in conversion of wood lignin: multicopper oxidases (MCOs, 18 genes) as laccases (Lcc), class II peroxidases (8 genes) as manganese peroxidases (MnP), glyoxal oxidases (5 genes, GLOX), quinone-reducing oxidoreductases (19 genes, QOR) and GMC oxidoreductases (12 genes) as aryl alcohol oxidases (AAO). We studied the genomic organisation and phylogeny of these genes as well as their expression using qRT-PCR. Comparative and phylogenetic analyses of genes coding for enzymes involved in wood lignin conversion and decomposition (i.e. lignin-modifying class II peroxidases) reveal differences between white- and brown-rot, necrotrophic and saprotrophic wood-decaying basidiomycetes. Transcript profiling using qRT-PCR revealed that some transcripts were very abundant in lignin-rich media, in cellulose-rich media, in wood or in the free-living mycelium grown liquid culture, suggesting specific functions of these genes, which need to be studied further.
Abstract
The causative agent of dieback on European ash (Fraxinus excelsior) was first described as Chalara fraxinea based on cultural morphology because no sexual stage of the fungus was known. Later, based on culturing of ascospores of a candidate teleomorph, morphological comparison and nuclear ribosomal internal transcribed spacer sequencing, the sexual stage of C. fraxinea was assigned as Hymenoscyphus albidus, a native and widespread species in Europe. Recently, the morphological species concept of H. albidus was shown to cover two species that cannot be separated from each other based on teleomorph characters, but which can be distinguished by several DNA markers. As a result, the strains causing ash dieback were reassigned as H. pseudoalbidus. The closely related H. albidus is presumably a non-pathogenic endophyte, but pathogenicity tests to confirm this hypothesis have not yet been performed. Genotyping of herbarium specimens has shown that H. pseudoalbidus was present in Switzerland for at least a decade prior to the epidemic outbreak in Europe. The origin of the ash dieback pathogen, and the general importance of correct pathogen identification to development of effective disease control, are discussed.
Authors
Tao Zhao Paal Krokene Jiang Hu Erik Christiansen Niklas Björklund Bo Långström Halvor Solheim Anna-Karin Borg-KarlsonAbstract
Background: Tree-killing bark beetles (Coleoptera, Scolytinae) are among the most economically and ecologically important forest pests in the northern hemisphere. Induction of terpenoid-based oleoresin has long been considered important in conifer defense against bark beetles, but it has been difficult to demonstrate a direct correlation between terpene levels and resistance to bark beetle colonization. Methods: To test for inhibitory effects of induced terpenes on colonization by the spruce bark beetle Ips typographus (L.) we inoculated 20 mature Norway spruce Picea abies (L.) Karsten trees with a virulent fungus associated with the beetle, Ceratocystis polonica (Siem.) C. Moreau, and investigated induced terpene levels and beetle colonization in the bark. Results: Fungal inoculation induced very strong and highly variable terpene accumulation 35 days after inoculation. Trees with high induced terpene levels (n = 7) had only 4.9% as many beetle attacks (5.1 vs. 103.5 attacks m22) and 2.6% as much gallery length (0.029 m m22 vs. 1.11 m m22) as trees with low terpene levels (n = 6). There was a highly significant rank correlation between terpene levels at day 35 and beetle colonization in individual trees. The relationship between induced terpene levels and beetle colonization was not linear but thresholded: above a low threshold concentration of ,100 mg terpene g21 dry phloem trees suffered only moderate beetle colonization, and above a high threshold of ,200 mg terpene g21 dry phloem trees were virtually unattacked. Conclusion/Significance: This is the first study demonstrating a dose-dependent relationship between induced terpenes and tree resistance to bark beetle colonization under field conditions, indicating that terpene induction may be instrumental in tree resistance. This knowledge could be useful for developing management strategies that decrease the impact of tree-killing bark beetles.
Abstract
No abstract has been registered
Abstract
Introduction: Windstorm is one of the most destructive environmental disturbance factors on forests, but its influence on conifer defense chemistry and susceptibility to insects and diseases is not well understood. Materials and methods: We selected groups of 10 Norway spruce trees with short leaders, leaning stems, or no apparent damage 17 months after the largest storm ever recorded in Sweden. Trees were mass-inoculated with Ceratocystis polonica, a virulent blue stain fungus associated with the spruce bark beetle (Ips typographus) to estimate tree resistance. Terpene and phenolic composition in the bark was analyzed by gas chromatography–mass spectrometry, two-dimensional gas chromatography, and liquid chromatography. Results: In contrast to our hypothesis, the results showed that trees with no apparent damage were more susceptible to C. polonica inoculation than short-leader and leaningstem trees. Chemical composition also differed between trees in different damage classes. (+)-3-carene and two unidentified stilbenes were higher, and taxifolin glycoside was lower in trees without apparent damage than in the others. The relative amount of (−)-α-pinene was negatively correlated, whereas (+)-3-carene, sabinene, (−)-germacrene D, thunbergol and two unidentified stilbenes were positively correlated with fungal performance. Conclusions: These results suggested that wind damage had increased resistance level of short-leader and leaning trees to C. polonica inoculation, and that change in terpene and phenolic composition in the bark could be at least partly responsible for the induced resistance. Different possible explanations for this unexpected finding are discussed.
Abstract
No abstract has been registered
Abstract
No abstract has been registered
Abstract
Plants are exposed to a variety of pathogens in their natural habitats. To understand the key processes of defense responses in aspen (Populus tremulae) at the transcript level two clones C72 and C23 with differential level of resistance from the SwAsp collection were inoculated with a foliar rust (Melampsora magnusiana Wagnar). Leaf samples were collected from adjacent areas of the inoculation site to examine the long distance (systemic) defense responses at day1, day3 and day14 post treatments. We performed microarray experiments on the biothrophic interaction, on comparison with the healthy controls we found that the two clones respond in a widely different fashion to the rust. Clone C23 showed almost no response to biotroph after 24 hours while clone 72 gave a clear defense response to the pathogen. Quantitative reverse-transcriptase polymerase chain reaction (qRT-PCR) showed a significant differential expression patterns in susceptible and resistant colnes. Chitinase, cinnamic acid reductase and the iaa genes showed signification up-regulation in resistant clone. The level of expression was 5.9 delta threshold cycles in chitinase gene at day14. Data analysis from extracted total phenolics and condensed tannins verify the results of cDNA arrays and qRT-PCR.
Abstract
Development in surface mould growth on painted/unpainted wooden claddings and acting climatic factors were investigated over a period of 3 years. Eight wood substrates, including modified, preservative-treated and untreated wood, were tested in combination with three types of paint: (1) water-borne alkyd modified acrylic paint without fungicide; (2) solvent-borne alkyd paint without fungicide; and (3) ICP (internal comparison product). One set of samples was exposed unpainted. The samples were tested according to a modified version of EN 927-3. A logistic regression model was fitted to the data. The degree of mould growth varied with exposure time, coating typology, wood substrate, temperature and relative humidity. Exposure time and coating typology contributed most to the model. After 3 years of outdoor exposure unpainted panels and panels coated with solvent-borne paint without fungicide had more mould growth than panels coated with ICP and water-borne paint without fungicide. Unpainted oil/copper–organic preservative-treated claddings had higher resistance to mould growth than other unpainted wood substrates. Coated untreated pine and coated acetylated pine were more susceptible to mould growth than other coated wooden substrates.
Authors
Igor A. Yakovlev Ari M. Hietala Pierre-Emmanuel Courty Francis Martin Jan Stenlid Halvor Solheim Carl Gunnar FossdalAbstract
The genome H. annosum s.l. was sequenced by JGI to a 8.23X coverage and assembled into 39 scaffolds with a total size of 33.7 Mb covering more than 98% of the whole genome. Based of genome sequence we annotated a number of genes for fungal enzymes that are believed to participate in lignin degradation, including: laccases (Lcc18 genes), manganese peroxidases (MnP8 genes) and hydrogen peroxide forming enzymes such as glyoxal oxidases (GLOX5 genes), quinone oxidoreductases (QOR17 genes) and aryl alcohol oxidases (AAO16 genes), which is in concordance with these gene family sizes observed in other sequenced white-rot fungi. We studied the genomic organisation and phylogeny of these genes as well as their expression using NimbleGen arrays and qRT-PCR. Transcript profiling using whole-genome oligo arrays and qRT-PCR revealed that some transcripts were very abundant in lignin-rich media (Lcc5 15, MnP2, GLOX4, QOR2 10, AAO9), in cellulose-rich media (lcc2, 7 16, MnP3 4, GLOX3, QOR4 6, AAO2, 7 10), in wood (Lcc3, MnP4, QOR2, GLOX1, AAO10) or in the free-living mycelium grown liquid culture (Lcc1, 3, 10 13), suggesting specific functions of these genes, which need to be studied further.
Abstract
Microbial disfigurement of coated wooden surfaces is considered a major maintenance concern and will shorten the aesthetic service life of wooden facades. The effect of the physical surface structure of a paint film when applied on wood may have an impact on the susceptibility to mould growth. Six model paints were formulated to give the following physical surface structures: glossy, matt, soft, hard, hydrophobic, and a film with air inclusion. The model paints and a standard paint, with and without fungicide, were applied on panels of Norway spruce (Picea abies L Karst.) and exposed outdoors for nearly three years according to a modified version of EN 927-3. A logistic regression model was fit to the data, and the degree of mould growth varied with exposure time and type of paint. Hard model paint was significantly more susceptible than the other model paints and had a performance close to the standard paint without fungicide. Soft model paint provided the best performance, with the least mould growth. Temperature, relative humidity, and precipitation did not significantly contribute to the model. (C) 2010 Elsevier Ltd. All rights reserved.
Authors
Gilbert Kamgan Nkuekam Halvor Solheim Z. Wilhelm De Beer Joha W. Grobbelaar Karin Jacobs Michael J. Wingfield Jolanda RouxAbstract
Ophiostoma spp. include important pathogens of trees and causal agents of sapstain. These fungi infect wounds on trees and are typically carried by insects, especially bark beetles. Ophiostoma spp. on coniferous hosts in the Northern Hemisphere are well-known. However, other than for the serious pathogens O. ulmi and O. novo-ulmi, very little research has been done on the occurrence of this group of fungi on native broad-leaved trees, especially in the Nordic countries. In this study, surveys were conducted in several areas of Norway to isolate Ophiostoma spp. associated with wounds on native broad-leaved trees belonging to the genera Betula, Fagus, Quercus, Sorbus and Tilia. Morphological studies and comparisons of DNA sequences for the ITS, 5.8S and part of the beta-tubulin gene regions were used to confirm the identity of the fungi collected. Ophiostoma spp., and especially their Pesotum anamorphs, were common on wounds on the trees sampled. In most cases, they were associated with wood stain. Ophiostoma spp. collected included predominantly O. quercus, O. borealis sp. nov., and O. denticiliatum. The results of this study emphasize that the diversity of Ophiostoma spp. on broad-leaved trees is still incompletely understood in Norway and other European countries.
Abstract
The blue-stain fungus Ceratocystis resinifera colonizes wounds on living Picea spp. and other conifers in Europe and North America. Little is known regarding the pathogenicity of this fungus and consequently, four Norwegian C. resinifera isolates were inoculated on to Norway spruce (Picea abies) using two different techniques. These included single-point inoculations on young trees (two inoculations per tree on 14-year-old trees) and mass-inoculations on older trees (∼200 inoculations per tree on 34-year-old trees). In both experiments, C. resinifera induced minor symptoms that in most cases did not differ significantly from inoculation with sterile agar. The virulent blue-stain fungus C. polonica, which was inoculated for comparative purposes, induced extensive symptoms, causing 83% dead cambium circumference and 82% blue-stained sapwood, and long necrotic lesions in the phloem. The results suggest that C. resinifera is non-pathogenic or only mildly pathogenic to Norway spruce and does not present a threat to these trees.
Authors
Leif Sundheim Trond Hofsvang Christer Magnusson Trond Rafoss Arild Sletten Halvor Solheim Arne Stensvand Brita Toppe Anne Marte Tronsmo Bjørn ØklandAbstract
No abstract has been registered
Authors
Leif Sundheim Trond Hofsvang Christer Magnusson Trond Rafoss Arild Sletten Halvor Solheim Brita Toppe Anne Marte Tronsmo Bjørn ØklandAbstract
No abstract has been registered
Authors
Leif Sundheim May Bente Brurberg Trond Hofsvang Christer Magnusson Trond Rafoss Arild Sletten Birger Solberg Halvor Solheim Brita Toppe Anne Marte Tronsmo Bjørn ØklandAbstract
No abstract has been registered
Authors
Tao Zhao Paal Krokene Niklas Björklund Bo Långström Halvor Solheim Erik Christiansen Anna-Karin Borg-KarlsonAbstract
Constitutive and inducible terpene production is involved in conifer resistance against bark beetles and their associated fungi. In this study 72 Norway spruce (Picea abies) were randomly assigned to methyl jasmonate (MJ) application, inoculation with the bluestain fungus Ceratocystis polonica, or no-treatment control. We investigated terpene levels in the stem bark of the trees before treatment, 30 days and one year after treatment using GC–MS and two-dimensional GC (2D-GC) with a chiral column, and monitored landing and attack rates of the spruce bark beetle, Ips typographus, on the trees by sticky traps and visual inspection. Thirty days after fungal inoculation the absolute amount and relative proportion of (+)-3-carene, sabinene, and terpinolene increased and (+)-α-pinene decreased. Spraying the stems with MJ tended to generally increase the concentration of most major terpenes with minor alteration to their relative proportions, but significant increases were only observed for (−)-β-pinene and (−)-limonene. Fungal inoculation significantly increased the enantiomeric ratio of (−)-α-pinene and (−)-limonene 1 month after treatment, whereas MJ only increased that of (−)-limonene. One year after treatment, both MJ and fungal inoculation increased the concentration of most terpenes relative to undisturbed control trees, with significant changes in (−)-β-pinene, (−)-β-phellandrene and some other compounds. Terpene levels did not change in untreated stem sections after treatment, and chemical induction by MJ and C. polonica thus seemed to be restricted to the treated stem section. The enantiomeric ratio of (−)-α-pinene was significantly higher and the relative proportions of (−)-limonene were significantly lower in trees that were attractive to bark beetles compared to unattractive trees. One month after fungal inoculation, the total amount of diterpenes was significantly higher in putative resistant trees with shorter lesion lengths than in putative susceptible trees with longer lesions. Thus, terpene composition in the stem bark may be related to resistance of Norway spruce against I. typographus and C. polonica.
Abstract
The anamorph genus Leptographium Lagerberg and Melin includes species that are typically bark beetleassociated fungi, with teleomorphs in Grosmannia. During a survey of ophiostomatoid fungi in Norway, two unusual species, that fit the broader morphological description of Leptographium, were isolated directly from the rootfeeding beetles, Dryocetes authographus and Hylastes cunicularius, as well as from roots infested by these insects. The first of these could be distinguished from other described species based on a sparse sporulation, black spore drops and chlamydospores in older cultures. This species also produces a Hyalorhinocladiella synanamorph. The second species was characterised by distinctly curved conidia. Based on these unusual morphological characteristics and distinct DNA sequences, these fungi were recognised as new taxa for which the names Leptographium chlamydatum sp. nov. and L. curvisporum sp. nov. are provided.
Abstract
European ash (Fraxinus excelsior), also known as common ash, occurs naturally inland in lower areas of southeastern Norway and along the southern coast of the country. It is important both as a forest and ornamental tree. During the last decade, dieback has become a disastrous disease on F. excelsior in many European countries. The anamorphic fungus Chalara fraxinea T. Kowalski (1), described for the first time from dying ash trees in Poland, is now considered the cause of ash dieback (2). In May of 2008, C. fraxinea was isolated from 1.5 m high diseased F. excelsior in a nursery in Østfold County in southeastern Norway. Symptoms included wilting, necrotic lesions around leaf scars and side branches, and discoloration of the wood. From symptomatic branches, small pieces (approximately 1 cm3) were excised in the transition area between healthy and discolored wood. After surface sterilization (10 s in 70% ethanol + 90 s in NaOCl), the pieces were air dried for 1 min in a safety cabinet, cut into smaller pieces, and placed on media. The fungus was isolated on potato dextrose agar (PDA) and water agar (WA). On PDA, the cultures were tomentose, light orange, and grew slowly (21 mm mean colony diameter after 2 weeks at room temperature). Typical morphological features of C. fraxinea developed in culture. Brownish phialides (14.8 to 30.0 [19.5] × 2.5 to 5.0 [4.1] μm, n = 50) first appeared in the center of the colonies on the agar plugs that had been transferred. The agar plugs were 21 days old when phialides were observed. Abundant sporulation occurred 3 days later. Conidia (phialospores) extruded apically from the phialides and formed droplets. Conidia measured 2.1 to 4.0 (3.0) × 1.4 to 1.9 (1.7) μm (n = 50). The first-formed conidia from each phialide were different in size and shape from the rest by being longer (6 μm, n = 10) and more narrow in the end that first appeared at the opening of the phialide. Internal transcribed spacer sequencing confirmed that the morphological identification was correct (Accession No. EU848544 in GenBank). A pathogenicity test was carried out in June of 2008 by carefully removing one leaf per plant on 10 to 25 cm high F. excelsior trees (18 trees) and placing agar plugs from a 31-day-old C. fraxinea culture (isolate number 10636) on the leaf scars and covering with Parafilm. After 46 days, isolations were carried out as described above from discolored wood that had developed underneath necrotic lesions in the bark and subsequently caused wilting of leaves. All the inoculated plants showed symptoms, and C. fraxinea was successfully reisolated. No symptoms were seen on uninoculated control plants (eight trees) that had received the same treatment except that sterile PDA agar plugs had been used.
Abstract
To identify differentially expressed genes of the white-rot fungus Heterobasidion parviporum subtractive cDNA libraries were constructed using suppressive subtraction hybridization (SSH) technique with RNA extracted from an advanced stage of decay area and from colonization front next to the reaction zone of the stem of a mature Norway spruce trees. Besides cytochrome P450s and proteins with unknown function, the SSH libraries constructed contained genes involved in basic cellular processes, andcell wall degradation. To examine the role of selected candidate genes three trees, showing a variable degree of wood decay, were used for real-time RT-PCR profiling of candidate genes. In the decay transition areas the study revealed activity centers that showed remarkable similarity in the transcript profiles of monitored genes.
Authors
Åke Olson Mikael Brandström Kajsa Himmelstrand Mårten Lind Magnus Karlsson K. Lunden Kerstin Dalman Heriberto Vélëz Björn Canbäck Francis Martin Halvor Solheim Carl Gunnar Fossdal Pierre Rouzé Yao-Cheng Lin Matteo Garbelotto Ursula Kües Igor V. Grigoriev Andrea Aerts Erika Lindquist Jeremy Schmutz Jan StenlidAbstract
The genome sequence of the conifer rot root pathogen Heterobasidion annosum was generated at JGI with 8.23 X coverage. The nuclear genome assembles in 39 scaffolds of total 33.7 Mbp estimated to cover 98.1% of the complete genome. We predicted 12,270 genes with an average length of 1,617 bp and exon number and length of 5.54 and 250 bp respectively. About 50% (5999) of the predicted genes could be validated by EST support with the 40,807 EST´s generated with in the project. The genome has a GC content of 52.0% and very little repetitive sequences with 2,895 SSR per mega base. The physical genome is congruent with the genetic linkage map, and most of the linkage groups have been possible to anchor to the 18 largest scaffolds.
Authors
Nadeem Yaqoob Igor A. Yakovlev Paal Krokene Harald Kvaalen Halvor Solheim Carl Gunnar FossdalAbstract
Clonal variation towards resistance has been observed in Norway spruce Heterobasidion annosum s.l. (H.a). H.a. is the main cause of root rot and has a severe economic impact on an economically important conifer tree species. Annual financial losses are in the hundreds of millions of Euros annually. Less susceptible clones appear to have an efficient system of recognizing the pathogen and initiating early defense signalling events. Active defense responses can be started locally and transmitted systemically. This work focus on the expression both spatially (systemically) and temporally in this pathosystem. Two-year-old, somatic saplings of the Norway spruce clone were challenged with H.a., wounded, methyl jasmonate painted and compared to untreated controls and ninety plants were used for the experiment. Stem samples were collected at 1, 3, 6 and 13 days post inoculation (d.p.i). The stem of the saplings were divided into sections along its length and the bark and wood separated from each other at time of collection. In order to see local response an area of 1cm including the site of inoculation was collected, while the spatial (systemic) response was assessed in sections collected at distances of 3 and 6cm away from the site of inoculation. The separated bark and wood were analysed for differential gene expression by qRT-PCR, and the results from peroxidases (PaPX3 and PaPX2) and a chitinase (PaChi4) are presented. Both local and systemic up- and down-regulation were observed at the transcriptional level in both bark and wood, up to 2000 fold local increase in expression was observed for PaChi4.
Abstract
Due to the great economic losses caused by the root and butt-rot pathogen Heterobasidion annosum, development of efficient control measures is warranted. H. annosum a necrotroph colonize the Norway spruce from inside and is responsible for 100s of millions of Euros losses annually. Considerable clonal variation has been recorded for Norway spruce in resistance towards H. annosum, but the defence mechanisms contributing to host resistance against both necrotrophic and biotrophic fungi remain poorly understood. The recent genome sequencing of Populus has made the genus a model to facilitate tree genetics. Genome-wide transcript profiling of Populus tremula upon pathogen attack will now be used, and homologues of Norway spruce genes to defence genes up-regulated in Populus will be identified. Two aspen clones (23 and 72) from the SwAsp collection (Luquez 2007) were used in the present study. Plants were propagated from tissue culture and kept in greenhouse under un-manipulated conditions. To study the host defence mechanisms, the rust (Melampsora magnusiana Wagner) and a bluestain fungus were used as a biotrophic and necrotrophic fungus respectively. Melampsora spores solution was applied to the underside of the leaf. To control for sectoriality six leaves were infected on each plant. To ensure high humidity and avoid cross contamination, plastic bags were wrapped around infected leaves. Leaves above infected leaves (10, 9, and 8) were harvested respectively 1, 3 and 14 days after inoculation. Initial results from microarray data indicate a clear separation between two Aspen clone (23 and 72) lines. For line 23 the response to biotroph and necrotroph seems to be similar. Whereas the response for line 72 is divergent for the treatments as they go in opposite direction. The controls show that there is an initial difference in the 2 lines (controls are separated). What are the genes that make biotrohic and necrotrophic treatment of 72 look so different will be worked out from microarray data. Differential expression of defense genes in biotrohic and necrotrophic treatment will be verified further with quantitative real time PCR. Chemical analysis of Aspen leaves gave less phenolic compounds as plants were kept in greenhouse. HPLC will be carried out after reaching some conclusion from microarray data analysis.
Abstract
Ophiostomatoid fungi were isolated from Scolytus ratzeburgi infesting Betula pendula and B. pubescens in Norway. Fungi were identified based on morphology, DNA sequence comparison for two gene regions and phylogenetic analyses. The most abundant fungus was Ophiostoma karelicum, suggesting a specific relationship between the fungus, the vector insect and the host tree. Our results suggest that O. karelicum occurs across the geographic range of S. ratzeburgi and its close relatedness to the Dutch elm disease fungi suggests that it could be important if introduced into other parts of the world. Other fungi, only occasionally isolated from S. ratzeburgi, were identified as O. quercus and a novel taxon, described here as O. denticiliatum sp. nov.
Authors
Ari Hietala Nina Elisabeth Nagy Arne Steffenrem Harald Kvaalen Carl Gunnar Fossdal Halvor SolheimAbstract
No abstract has been registered
Abstract
Two mature clones of Norway spruce (Picea abies (L.) Karst.) shown to have different level of resistance towards inoculation of Heterobasidion parviporum were compared with respect to spatiotemporal expression of transcripts related to biosynthesis of lignin, stilbenes and other phenolic compounds in response to fungal inoculation and physical wounding. Both clones responded to H. parviporum and physical wounding at transcriptional and chemical levels. Taxifolin, detected in the resistant clone only, increased in concentration following both wounding and inoculation. Concentrations of stilbenoid glucosides were highest in the susceptible clone. Following wounding or inoculation, concentrations of these glucosides increased in the susceptible clone, and quantities of their corresponding aglycones increased dramatically in both clones close to the treatment point. Significant changes in transcription were detected over the entire lesion length for all transcripts, and only the changes in a few transcripts indicated a response to inoculation with H. parviporum differing from that caused by wounding alone. The resistant clone had higher basal concentrations of lignin (LTGA) compared to the susceptible clone; concentrations increased in both clones after wounding and wounding plus inoculation treatments, but remained consistently higher in the resistant clone, suggesting higher lignin levels in the cell walls compared to the susceptible clone. In addition, the transcript level in the same clones was also measured the following year and we saw indications of primed defences for a number of gene products likely resulting from the inoculations performed 12 months prior.
Abstract
The root-rot causing fungus Heterobasidion annosum senso lato is the most devastating pathogen of conifers in Europe. This pathogen enter Norway spruce trees trough the roots and colonizes the tree from within, growing as a saprophyte when established within the dead heartwood and acting as a necrotroph when in contact with living host tissue. The genome of this devastating pathogen has now been sequenced in collaboration with JGI and gene annotation is ongoing and genomic work is currently in progress (Stenlid et al. work in progress). We have worked with the host Norway spruce from a molecular perspective for more than ten years. Twenty percent of the trees in Norwegian spruce stands tend to be infected and this pathogen that can colonize ten meters up inside the trunk. The tree have defences against this pathogen and the attack can be fought off by the bark and living wood but not by the hearthwood. The tree has a unique defense against this internal attack by forming a reaction zone; in this case the host defense is directed inwardly by the still living sapwood toward the central colonized wood. We have in the last years studied the host responses to infection in Norway spruce clones at the transcriptional level and found that the speed of recognition and spatial defense signalling appears to be the hallmarks of trees with high degree of resistance. We strive to study both partners in this pathosystem from a molecular perspective, and are using suppressive subtractive hybridization (SSH) followed by Real-Time RT PCR verification to look at differentially expressed genes(Yakovlev et al. 2008). In addition the colonization profiles are followed on extracted gDNA using quantitative Real-Time PCR (Hietala et al. 2009).
Abstract
In natural conditions plants are continuously exposed to number of pathogens both biotrophs and necrotrophs. To understand their defense response at the transcript level two clones C72 and C23 with differential level of resistance from the SwAsp collection were inoculated with a biotroph (Melampsora magnusiana Wagnar) and necrotroph (Ceratocysis spp.) and compared to wounded and healthy controls. Samples were collected in leaves and areas some distance away from the inoculation site to examine the long distance (systemic) defense responses at day, day3 and day14 post treatments. We performed microarray experiments on the necrotrophic and biothrophic interaction compared with the healthy controls and found that the two clones respond in widely different fashions to the treatments applied. Clone C23 showed almost no response to biotroph and necrotroph inoculations after 24 hours while clone 72 gave a clear defense response to both pathogens. We are now in the process of verifying these results and looking at additional time-points using qRT-PCR.
Abstract
The inhibitory effect of methanol bark extracts from six deciduous and three coniferous European tree species were bioassayed against eight fungi from the different damage categories, brown rot, white rot, canker and blue-stain. This is the first report providing data on the antifungal activity of several Europaen tree species against fungi within these damage categories. Generally the decay fungi were more inhibited by the bark extracts than the blue-stain fungi, while the lowest inhibition was found among the cancer fungi. The main pattern found between the fungal groups in relation to the bark extracts in this study is believed to be caused by the route of ingress. Acer platanoides bark extract proved to be the most effcient bark extract tested, significantly reducing the growth rate of all tested fungi. Betula pubescens bark extract generally gave the weakest reduction in growth rate. In this study, the conifer bark extracts were in general more active against the canker and blue stain ascomycete fungi than the deciduous trees extracts.
Abstract
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Authors
Halvor SolheimAbstract
Climate change may influence in a worse manner for the forests in various ways. Some pathogens may increase their importance and new may arrive. Root and butt rot is the most serious problem in Norway spruce forests. In mean more than every fourth tree is infested when harvested. Dryer summers may give increased frequency of rot caused by Heterobasidion. In addition Armillaria spp may gain change in weather condition both as root rot and in connection with a syndrome together with drought and bark beetles. More unstable winter climate may give increase of Gremmeniella attack on Scots pine. Longer and warmer growth season will give many pathogens better condition. Among those is Ophiostoma novo-ulmi causing Dutch elm disease which is lasting in south eastern Norway at a rather low frequency and the volume of elm is not lower than for 15 years ago. In which way the newly introduced Chalara fraxinea will behave in Norway is uncertain, but a better growth season will probably also influence on the possibility to be spread all over Norway where ash are growing.
Abstract
Ips typographus is economically most important insect pest of mature spruce in Eurasia. Normally, it prefers to reproduce in dead and/or dying trees, but following large-scale storm disturbances, its outbreaks kill waste areas of living stands. One factor triggering such epidemics is a surplus of broken and uprooted trees with non-existent, or weak, defence....
Abstract
Induced reactions in the phloem is a basic mechanism of conifer resistance to bark beetle and their associated fungi (1,2). Previous research has proved that certain doses of Ceratocystis polonica infection or methyl jasmonate (MeJA) application could induce acquired resistance and decrease subsequent fungal or bark beetle colonization (3,4,5). To study the induced chemical changes after fungal infection and MeJA application in the phloem of mature Norway spruce, three groups, each of 24 P. abies trees of similar size, were chosen in Tönnersjöheden, southern Sweden, in May 2006. The three groups were then inoculated with C. polonica, sprayed with MeJA, or used as untreated control, respectively. Phloem samples were taken twice from each tree: on the same day as treatment and 1 mo later. The terpene composition of all the samples was analyzed by GC-MS, and the enantiomeric compositions of α-pinene, β-pinene, and limonene were analyzed by 2D-GC (6). The result indicated that both MeJA application and C. polonica infection had certain effects on the terpene composition. C. polonica infection significantly increased the biosynthesis of 3-carene, sabinene, and terpinolene. Both mean absolute amounts and relative amounts of these monoterpenes increased in samples from fungus inoculated trees, similar to what is observed in Scots pine after Leptographium wingfieldii inoculation (7). MeJA application increased the absolute amount of α-pinene, β-pinene, limonene, and some other major terpenes, but it did not change the relative amount of these terpenes. However, neither MeJA application nor fungal infection changed the enantiomeric compositions of α-pinene, β-pinene, and limonene in the phloem of Norway spruce.
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Abstract
The root-rot causing fungus Heterobasidion annosum sensu lato is the most devastating pathogen of conifers in Europe. This pathogen enters Norway spruce through the roots and can colonize the tree from within, growing as a saprophyte when established within the dead heartwood and acting as a necrotroph when in contact with living host tissue. Despite the high incidence of damage, trees have defences against this pathogen in the bark and living wood. Furthermore, spruce has a defense against internal attack by forming a reaction zone, in this case the host defense is directed inwardly by the still living sapwood toward the central colonized heartwood. We have studied the host responses to infection in Norway spruce clones at the transcriptional level and found that the speed of recognition and that spatial defense signalling appears to be the hallmarks of trees with high degree of resistance...
Abstract
The difficulty in subculturing biotrophic fungi complicates etiological studies related to the associated plant diseases. By employing internal transcribed spacer rDNA-targeted quantitative real-time polymerase chain reaction, we now show that the heteroecious rust Thekopsora areolata, commonly associated in natural conditions to sapling shoots and cones of Norway spruce and leaves of wild bird cherry, frequently infects nurserygrown seedlings of the conifer. A spatial sampling scheme was used to investigate seedlings and saplings of Norway spruce showing phloem necrosis: the highest concentration of DNA of T. areolata was recorded in the area with necrotic phloem. The separate analysis of bark and wood tissues suggested that the initial spread of the rust to healthy tissues neighboring the infection site takes place in the bark. A Phomopsis species found to coexist with T. areolata in several seedlings showed very high DNA levels in the upper part of the lesion, and even in the visually healthy proximal tissues above the lesions, which indicates that the ascomycete, most probably a secondary invader following primary infection by T. areolata, has a latent stage during early host colonization. We hypothesize that this hemibiotrophic mode of infection contributes to the successful coexistence of Phomopsis with a biotrophic rust.
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Abstract
Due to the great economic losses caused by the root and butt-rot pathogen Heterobasidion annosum, developments of efficient control measures are warranted. H. annosum a necrotroph colonize the Norway spruce from inside and is responsible of 10-13 millions Euros losses in Norway alone. Considerable clonal variation has been recorded for Norway spruce in resistance towards H. annosum, but the defence mechanisms contributing to host resistance remain poorly understood. The recent genome sequencing of Populus has made the genus a model to facilitate tree genetics. Genome-wide transcript profiling of Populus tremula upon pathogen attack will now be used, and homologues of Norway spruce genes to defence genes up-regulated in Populus will be identified. Populus-Phellinus tremula pathosystem is selected as P. tremula behaves like H. annosum.
Abstract
The Norwegian Monitoring Programme for Forest Damage (OPS) has since its start registered damage to selected trees. The aim of the registrations has been to explain variations in crown density and crown colour. In answer to international requests, the Norwegian Forest and Landscape Institute has prepared a short guide to the determination of the most common forms of damage found in Norwegian forests...
Abstract
The root-rot causing fungus Heterobasidion annosum senso lato is the most devastating pathogen of conifers in Europe. This pathogen enter Norway spruce trees trough the roots and colonizes the tree from within, growing as a saprophyte when established within the dead heartwood and acting as a necrotroph when in contact with living host tissue. Twenty percent of the trees in Norwegian spruce stands tend to be infected and this pathogen that can colonize ten meters up inside the tree trunk, decaying the silvicultural most valuable part of the tree. Despite this high incidence of damage the tree has efficient defences against this pathogen and the attack is eventually fought off if present in the bark or living wood. The tree also has a defense against this internal attack (by Heterobasidion established in the heartwood expanding and invading outward toward the living sapwood) by forming a reaction zone; in this case the host defense is directed inwardly by the still living sapwood toward the central colonized wood. We have in the last years studied the host responses to infection in Norway spruce clones at the transcriptional level and found that the speed of recognition and that spatial defense signalling appears to be the hallmarks of trees with high degree of resistance. We strive to study both partners in this pathosystem from a molecular perspective, and are now focusing on the pathogen and what fungal gene-products are being expressed during the colonization of the heartwood compared to those expressed close to the active host defense (reaction zone) using suppressive subtractive hybridization (SSH) followed by Real-Time RT PCR analysis. In addition the colonization profiles were followed on extracted gDNA using quantitative Real-Time PCR.
Authors
Nicolette Fouche Jolanda Roux Halvor Solheim Ronald N. Heath Gilbert Kamgan Nkuekam P. Chimwamurumbe K.G. Pegg Michael J. WingfieldAbstract
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Abstract
The area of wood protection is in a period of change. New tools are needed to understand the mode of action, and to further improve the new wood protection systems. A set of useful tools are found among the molecular methods. This paper presents an overview of some of the tools available, and the methods are exemplified by papers within the frame of wood protection issues. However, there is still a great unexplored potential within the field of wood protection by the use of various molecular methods. The majority of the work using molecular methods has been performed on species identification issues and within species variation. This paper lists some new promising molecular methods for wood protection issues and a presentation of a new project. The new project will help to gain some new knowledge about how the fungal decay processes are affected by different wood modification systems.
Authors
Karlsson Magnus Ari Hietala Harald Kvaalen Halvor Solheim Åke Olson Jan Stenlid Carl Gunnar FossdalAbstract
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Abstract
In winter 2000-2001, there was a serious outbreak of Gremmeniella abietina Morelet in southeastern Norway. During the outbreak, we noted that injured Scots pine trees (Pinus sylvestris L.) developed secondary buds in response to the fungus attack, and we decided to study the relationship between injury, appearance of secondary buds and recovery of the trees thereafter. For this purpose, 143 trees from 10 to 50 years of age were chosen and grouped into crown density classes. Injury was assessed in detail, and buds were counted before bud burst in the spring of 2002. In addition, a subset of 15 trees was followed through the summer of 2002 to assess recovery. All injured trees developed secondary buds, with a clear overweight of dormant winter buds in proportion to interfoliar buds. Healthy control trees did not develop secondary buds at all. The secondary buds appeared predominantly on the injured parts of the tree; interfoliar buds in particular developed just beneath the damaged tissue. Most of the secondary buds died during the winter of 2001-2002, mainly because the fungus continued to spread after the first outbreak. Many of the remaining buds developed shoots with abnormal growth during the summer. Secondary buds may help trees to recover from Gremmeniella attacks, but this strategy may fail when the fungus continues to grow and injure the newly formed buds and shoots.
Abstract
In spring 2002, extensive damage was recorded in southeast Norway on nursery-grown Norway spruce seedlings that had either wintered in nursery cold storage or had been planted out in autumn 2001. The damage was characterised by a top shoot dieback. Two visually distinct types of necroses were located either on the upper or lower part of the 2001-year-shoot. Isolations from the upper stem necroses rendered Gremmeniella abietina, while Phomopsis sp. was isolated mostly from the from the lower stem necroses. RAMS (random amplified microsatellites) profiling indicated that the G. abietina strains associated with diseased nursery seedlings belonged to LTT (large-tree type) ecotype, and inoculation tests confirmed their pathogenicity on Norway spruce seedlings. Phomopsis sp. was not pathogenic in inoculation tests, this implying it may be a secondary colonizer. We describe here the Gremmeniella - associated shoot dieback symptoms on Norway spruce seedlings and conclude that the unusual disease outburst was related to the Gremmeniella epidemic caused by the LTT type on large pines in 2001. The role of Phomopsis sp. in the tissue of diseased Norway spruce seedlings is yet unclear.
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The difficulty in sub-culturing biotrophic fungi complicates etiological studies related to the associated plant diseases. By employing species-specific ITS sequence stretches, we used real-time PCR to investigate the spatial colonization profiles of T. areolata and a co-existing Phomopsis species in seedlings and saplings of Norway spruce showing bark necrosis. There was a strong gradient in the colonization level of T. areolata DNA along the lesion length, with the highest DNA amount levels being recorded in the area with dark brown phloem. The separate analysis of bark and wood tissues indicated that the initial spread of the rust to healthy tissues neighbouring the infection site presumably takes place in the bark. A Phomopsis species co-existing together with T. areolata in several cases showed very high DNA levels in the upper part of the lesion outside the brown phloem area, and even in the visually healthy proximal tissues above the lesions. This indicates that this ascomycete has a latent stage during early colonization of Norway spruce shoots. This mode of infection most probably explains the successful co-existence of Phomopsis with a biotrophic rust, as their mutual interest would be to avoid triggering host cell death.
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Abstract
The root-rot causing fungus Heterobasidion annosum can attack both spruce and pine trees and is the economically most damaging pathogen in northern European forestry. We have monitored the H. annosum S-type (fairly recently named H. parviporum) colonization rate and expression of host chitinases and other host transcripts in Norway spruce material with differing resistances using quatitative realtime PCR. Transcript levels of three chitinases, representing classes I, II and IV, were monitored. Ramets of two 33-year-old clones differing in resistance were employed as host material and inoculation and wounding was performed. clones in the area immediately adjacent to inoculation. Fourteen days after infection, pathogen colonization was restricted to the area immediately adjacent to the site of inoculation for the strong clone (589), but had progressed further into the host tissue in the weak clone (409). Transcript levels of the class II and IV chitinases increased following wounding or inoculation, while the transcript level of the class I chitinase declined following these treatments. Transcript levels of the class II and class IV chitinases were higher in areas immediately adjacent to the inoculation site in 589 than in similar sites in 409 three days after inoculation, suggesting that the clones differ in the rate of pathogen perception and host defense signal transduction. This an earlier experiments using mature spruce clones as substrate indicate that it is the speed of the host response and not maximum amplitude of the host response that is the most crucial component in an efficient defense in Norway spruce toward pathogenic fungi such as H. annosum.
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Twenty-five tree species were recorded as hosts for five European Armillaria species in studies on forest ecosystems in Serbia. Armillaria was most frequently isolated from the conifers Picea abies and Abies alba and from the deciduous trees Fagus moesiaca and Quercus petraea. A. mellea and A. gallica coexisted in hardwood forests in northern and central parts of Serbia, while A. ostoyae and A. cepistipes were mostly present in coniferous forests in the southern mountain region of Serbia. The distribution depended on the Armillaria species, altitude, and the forest type.
Abstract
To identify chemical resistant markers induced by fungal or mechanical injury, young trees of Scots pine (Pinus sylvestris) were subjected to inoculations of blue stain fungi associated with the pine shoot beetles Tomicus piniperda and T. minor. Among the 20 trees selected for chemical analyses, 16 were divided into four groups: one as control and three were pretreated by wounding only, or by inoculation with either the blue stain fungus Leptographium wingfieldii or Ophiostoma canum.Four wk after pretreatment, all 16 pretreated trees were mass-inoculated with L. wingfieldii. The absolute and relative amounts, as well as the enantiomeric compositions of monoterpene hydrocarbons in the phloem, were determined via a small sample of the phloem before and after the pretreatment and mass inoculation, by using two-dimensional gas chromatography (2D GC) and GC-mass spectrometry (MS).After mass inoculation, the absolute amounts of most of the monoterpenes decreased in the phloem sampled 20 cm from the fungal infection, and were higher in the phloem sampled within the infected reaction zone.The relative amounts of both ()--pinene and ()-limonene increased in phloem samples taken 20 cm above the fungal inoculation in the preinoculated trees compared with phloem sampled from the remaining four control trees. The enantiomeric compositions of -pinene and limonene changed, after fungal growth, at defined distances from the inoculation site: the proportion of the ()-enantiomers was highest in the phloem sampled 20 cm from the fungal inoculation.Four wk after pretreatment, monoterpene production in the phloem at the site of inoculation was more enhanced by L. wingfieldii than by O. canum. However, the different virulence levels of the fungi did not affect the enantiomeric composition of the monoterpenes. The biosynthesis of monoterpene enantiomers is discussed in relation to induced pathogen resistance.
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Authors
Magnus Karlsson Ari M. Hietala Harald Kvaalen Halvor Solheim Åke Olson Jan Stenlid Carl Gunnar FossdalAbstract
Norway spruce (Picea abies (L.) Karst.) has a natural distribution in the northern parts of Europe and Asia and is economically the most important tree species grown in the Nordic countries. A common threat to Norway spruce is the basidiomyceteous fungus Heterobasidion parviporum Niemelä and Korhonen. H. parviporum mainly attacks Norway spruce, although Siberian fir (Abies sibirica Ledeb.) and Scots pine (Pinus sylvestris L.) occasionally get infected. One obstacle to studying host/pathogen interaction in conifers has been the limited availability of mature clones for controlled inoculations, as genetic variation within the host material and the lack of replicates complicate interpretation of the results. Somatic embryogenesis, rooted cuttings, and tissue cultures may provide solutions for this problem. Tissue cultures from mature Norway spruce trees have been proposed as a possible model system for assessing resistance toward fungal pathogens. Recent data on chitinase isoform activity in the Norway spruce/H. parviporum pathosystem are encouraging; clonal variation was observed in the isoforms affected by inoculation, and the isoforms showing increased band intensity following bark inoculation by H. parviporum were also induced in the inoculated tissue cultures of the corresponding clones. To investigate the biological relevance of tissue cultures in host-pathogen interaction studies, transcript levels of selected host and pathogen genes in tissue cultures of Norway spruce were compared to those in bark of 33-year-old ramets of the same clones upon challenge by the pathogenic fungus H. parviporum. Similar transcript profiles of the pathogen and host genes were observed in both tissues, this supporting the use of tissue cultures as experimental material for the pathosystem. Higher transcript levels of the host genes phenylalanine ammonia lyase, peroxidase, and glutathione-S-transferase were observed in the more resistant clone #589 than in the less resistant clone #409 during the early stages of colonization. The most striking difference between the spruce clones was related to gene transcript levels of a class IV chitinase, which showed a continuous increase in clone #409 over the experimental period, with a possible association of this gene product to programmed cell death. Several of the fungal genes assayed were differentially expressed during colonization, including putative glutathione-S-transferases, laccase, cellulase, cytochrome P450 and superoxide dismutase genes. The transcriptional responses suggest an important role for the antioxidant systems of both organisms.
Abstract
In spring 2002, extensive damage was recorded in southeast Norway on nursery-grown Norway spruce seedlings that had either wintered in nursery cold storage or had been planted out in autumn 2001. The damage was characterised by a top shoot dieback. Two visually distinct types of necroses were located either on the upper or lower part of the 2001-year-shoot. Isolations from the upper stem necroses rendered Gremmeniella abietina, while Phomopsis sp. was isolated mostly from the lower stem necroses. RAMS (random amplified microsatellites) profiling indicated that the G. abietina strains associated with diseased nursery seedlings belonged to LTT (large-tree type) ecotype,and inoculation tests confirmed their pathogenicity on Norway spruce seedlings. Phomopsis sp. was not pathogenic in inoculation tests, this implying it may be a secondary colonizer. We describe here the Gremmeniella – associated shoot dieback symptoms on Norway spruce seedlings and conclude that the unusual disease outburst was related to the Gremmeniella epidemic caused by the LTT ecotype on large Scots pines in 2001. The role of Phomopsis sp. in the tissue of diseased Norway spruce seedlings is yet unclear.
Authors
Halvor SolheimAbstract
Norway spruce suffers from serious root and butt rot problems from sea level up to the timber line in Norway. In this paper the most common fungi causing white rot is presented with special notes on gross characteristics of the rot. During the meeting we visited a stand near the timberline where logging was ongoing. Isolations were done from nearly hundred rotten logs and the results are presented.
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In field experiments, clones of Norway spruce [Picea abies (L.) Karst.] showed different degrees of resistance against pathogenic fungi inoculated into the bark that correlate with differences in polyphenolic parenchyma (PP) cells of the bark. Cells of spruce callus cultures, particularly towards the callus surface, resemble PP cells and this study looks at changes in callus cells during infection and the relative resistance of cultures from clones of low (weak) or high (strong) resistance to fungal infection. Callus cultures, initiated from trees with different resistance, were co-inoculated with Ceratocystis polonica (Siem.) C. Moreau and Heterobasidion annosum (Fr.) Bref. Callus cells from strong clones resemble PP cells of bark tissue from strong clones, having more polyphenolic bodies, while callus cells from weak clones are more similar to PP cells from those clones, which have less extensive phenolic bodies. Callus cultures from trees with weak resistance were more quickly overgrown by both species of pathogenic fungi than cultures from trees with strong resistance. Callus cells of infected cultures showed changes similar to activated PP cells of bark, including enhanced accumulation of polyphenolics. Phenolic bodies were more numerous and more extensive (larger and denser) in callus cells of strong versus weak clones under all conditions. Thus, callus cells may perform similar functions in defense as PP cells in the bark. Callus from trees of varying resistance seem to reflect the relative resistance of the trees from which they are derived, and this study indicates that some mechanisms of resistance can be studied using callus from trees of different resistance.