Strigolactones were originally identified as stimulators of the germination of root-parasitic weeds 1 that pose a serious threat to resource-limited agriculture 2 . They are mostly exuded from roots and function as signalling compounds in the initiation of arbuscular mycorrhizae 3 , which are plant-fungus symbionts with a global effect on carbon and phosphate cycling 4 . Recently, strigolactones were established to be phytohormones that regulate plant shoot architecture by inhibiting the outgrowth of axillary buds 5,6 . Despite their importance, it is not known how strigolactones are transported. ATP-binding cassette (ABC) transporters, however, are known to have functions in phytohormone translocation [7][8][9] . Here we show that the Petunia hybrida ABC transporter PDR1 has a key role in regulating the development of arbuscular mycorrhizae and axillary branches, by functioning as a cellular strigolactone exporter. P. hybrida pdr1 mutants are defective in strigolactone exudation from their roots, resulting in reduced symbiotic interactions. Above ground, pdr1 mutants have an enhanced branching phenotype, which is indicative of impaired strigolactone allocation. Overexpression of Petunia axillaris PDR1 in Arabidopsis thaliana results in increased tolerance to high concentrations of a synthetic strigolactone, consistent with increased export of strigolactones from the roots. PDR1 is the first known component in strigolactone transport, providing new opportunities for investigating and manipulating strigolactone-dependent processes.Strigolactones are a new class of carotenoid-derived 10 phytohormone in land plants. In addition to their role in shoot branching, strigolactones are exuded into the rhizosphere under phosphorus-limiting conditions 5 and act as growth stimulants of arbuscular mycorrhizal fungi 3 . To identify efflux carriers of arbuscular-mycorrhiza-promoting factors such as strigolactones, we used a degenerate primer approach ( Supplementary Fig. 2a) to isolate full-size PDR-type transporters (also known as ABC subtype G (ABCG) transporters) of P. hybrida that are abundant in phosphate-starved or mycorrhizal roots. The rationale behind the focus on these transporters, of which there are 15 in Arabidopsis 11 , 23 in Oryza sativa (rice) 11 and 23 putative factors in Solanum lycopersicum (tomato) ( Supplementary Fig. 3a), was that they are plasma membrane proteins often found in roots 12 , they are implicated in below-ground plantmicrobe interactions 13,14 , and they have affinities for compounds that are structurally related to strigolactones 8,9,15 . Of six primary candidates, only P. hybrida PDR1 had increased expression in roots that were subjected to either phosphate starvation (Fig. 1a) or colonization by the arbuscular mycorrhizal fungus Glomus intraradices (Fig. 1b). Furthermore, PDR1 transcript levels increased in response to treatment with the synthetic strigolactone analogue GR24 or the auxin analogue 1-naphthaleneacetic acid (NAA) (Fig. 1c). Auxin has been shown to upregulate strigolactone-bi...

Forest pathology, the science of forest health and tree diseases, is operating in a rapidly developing environment. Most importantly, global trade and climate change are increasing the threat to forest ecosystems posed by new diseases. Various studies relevant to forest pathology in a changing world are accumulating, thus making it necessary to provide an update of recent literature. In this contribution, we summarize research at the interface between forest pathology and landscape ecology, biogeography, global change science and research on tree endophytes. Regional outbreaks of tree diseases are requiring interdisciplinary collaboration, e.g. between forest pathologists and landscape ecologists. When tree pathogens are widely distributed, the factors determining their broad-scale distribution can be studied using a biogeographic approach. Global change, the combination of climate and land use change, increased pollution, trade and urbanization, as well as invasive species, will influence the effects of forest disturbances such as wildfires, droughts, storms, diseases and insect outbreaks, thus affecting the health and resilience of forest ecosystems worldwide. Tree endophytes can contribute to biological control of infectious diseases, enhance tolerance to environmental stress or behave as opportunistic weak pathogens potentially competing with more harmful ones. New molecular techniques are available for studying the complete tree endobiome under the influence of global change stressors from the landscape to the intercontinental level. Given that exotic tree diseases have both ecologic and economic consequences, we call for increased interdisciplinary collaboration in the coming decades between forest pathologists and researchers studying endophytes with tree geneticists, evolutionary and landscape ecologists, biogeographers, conservation biologists and global change scientists and outline interdisciplinary research gaps.

BackgroundWhereas an increasing number of pathogenic and mutualistic ascomycetous species were sequenced in the past decade, species showing a seemingly neutral association such as root endophytes received less attention. In the present study, the genome of Phialocephala subalpina, the most frequent species of the Phialocephala fortinii s.l. – Acephala applanata species complex, was sequenced for insight in the genome structure and gene inventory of these wide-spread root endophytes.ResultsThe genome of P. subalpina was sequenced using Roche/454 GS FLX technology and a whole genome shotgun strategy. The assembly resulted in 205 scaffolds and a genome size of 69.7 Mb. The expanded genome size in P. subalpina was not due to the proliferation of transposable elements or other repeats, as is the case with other ascomycetous genomes. Instead, P. subalpina revealed an expanded gene inventory that includes 20,173 gene models. Comparative genome analysis of P. subalpina with 13 ascomycetes shows that P. subalpina uses a versatile gene inventory including genes specific for pathogens and saprophytes. Moreover, the gene inventory for carbohydrate active enzymes (CAZymes) was expanded including genes involved in degradation of biopolymers, such as pectin, hemicellulose, cellulose and lignin.ConclusionsThe analysis of a globally distributed root endophyte allowed detailed insights in the gene inventory and genome organization of a yet largely neglected group of organisms. We showed that the ubiquitous root endophyte P. subalpina has a broad gene inventory that links pathogenic and saprophytic lifestyles.Electronic supplementary materialThe online version of this article (doi:10.1186/s12864-016-3369-8) contains supplementary material, which is available to authorized users.

While Hymenoscyphus fraxineus causes dieback of the European ash (Fraxinus excelsior), flowering ash (F. ornus) appears resistant to the pathogen. To date, contributions of endophytic fungi to host resistance are unknown. The following hypotheses were tested: (i) endophytic fungi enhance the resistance of F. excelsior to the pathogen; (ii) resistance of F. ornus relies on its community of endophytic fungi. Two experiments were performed. (i) The effect of exudates of ash endophytes on the germination rate of H. fraxineus ascospores was studied in vitro Isolates of abundant Fraxinus leaf endophytes, such as Venturia fraxini, Paraconiothyrium sp., Boeremia exigua, Kretzschmaria deusta and Neofabraea alba inhibited ascospore germination. (ii) Ash seedlings inoculated in a climate chamber, with fungi sporulating on the previous year's leaf litter, were exposed to natural infections by the pathogen present in the forest. Non-inoculated seedlings were used as controls. Venturia spp. dominated the inoculated endophyte 'communities'. Subsequent exposure to H. fraxineus led to infection of F. excelsior leaves by the pathogen, but no differences in health status between pre-inoculated and non-inoculated seedlings were detected. Fraxinus ornus leaves experienced a low infection rate, independent of their colonization by endophytic fungi. These results did not support either hypothesis.

The European ash (Fraxinus excelsior) is threatened by the introduced ascomycete Hymenoscyphus fraxineus, the causal agent of ash dieback. Endophytic fungi are known to modulate their host’s resistance against pathogens. To understand possible consequences of ash dieback on the endophytic mycobiome, F. excelsior leaves were collected in naturally regenerated forests and the fungal communities analyzed by classic culture and Illumina amplicon sequencing using a newly developed and validated fungal-specific primer. Collections were done in the area infested by ash dieback north of the Alps, and in the disease free area on the south side. Sycamore maple (Acer pseudoplatanus) was additionally collected, as well as the flowering ash (F. ornus), which occurs naturally in the south and shows tolerance to ash dieback. Both cultivation and amplicon sequencing revealed characteristic endophytic fungal communities dominated by several strictly host specific Venturia species. On A. pseudoplatanus, a hitherto undescribed Venturia species was identified. Due to its dominance on F. excelsior, V. fraxini is unlikely to go extinct in case of reduced host densities. A majority of species was not strictly host specific and is therefore likely less affected by ash dieback in the future. Still, shifts in community structure and loss of genetic diversity cannot be excluded. The potentially endangered endophyte Hymenoscyphus albidus was rarely found. In addition to host specificity, species with preferences for leaf laminae or petioles were found. We also detected considerable geographical variation between sampling sites and clear differences between the two sides of the Alps for endophytes of F. excelsior, but not A. pseudoplatanus. Since sycamore maple is not affected by an epidemic, this could point toward an influence of ash dieback on ash communities, although firm conclusions are not possible because of host preferences and climatic differences. Furthermore, the mycobiota of F. excelsior trees with or without dieback symptoms were compared, but no clear differences were detected. Besides methodical refinement, our study provides comprehensive data on the ash mycobiome that we expect to be subject to changes caused by an emerging disease of the host tree.