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Skrede, I.

Publications and source records attributed to Skrede, I..

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Phenotypic Variation Reveals Contrasting Ecological Strategies in Wood-Decay Fungal Ecotypes Across a Hybrid Zone

Fungi play central roles in terrestrial ecosystem functioning and are the major decomposers of dead wood in forest systems. Wood decay fungi are adapted to growth and decay under different environmental conditions, but we have limited insight into the intraspecific variability in fungal growth and decay. In Fennoscandia, there are two genetically and ecologically distinct ecotypes of the wood-decay fungus Meruliopsis taxicola: a northern Continental ecotype associated with Norway spruce (Picea abies) growing in moist old-growth forests, and a southern Coastal ecotype growing on Scots pine (Pinus sylvestris) in harsher habitats. The two ecotypes hybridize in a narrow contact zone running through Fennoscandia. Here, we investigate the level of adaptation the two ecotypes show in phenotypic traits, and how hybrid isolates perform as compared with the parental genotypes. We performed in vitro experiments to quantify mycelial growth rate under varying temperature and drought conditions, as well as decomposition of the two substrates, Scots pine and Norway spruce. Isolates of the Continental ecotype exhibited generally higher growth rates in all environments and caused higher mass loss of both substrates. This is consistent with a more competitive life history strategy in the Continental ecotype, whereas Coastal isolates showed adaptions indicative of greater stress tolerance. Hybrid isolates displayed largely intermediate growth responses relative to the parental ecotypes. Together, these results reveal clear phenotypic divergence between M. taxicola ecotypes consistent with contrasting life-history strategies.

evolutionary biology↗

Long-term shift in community composition of deadwood fungi after clear-cutting

O_LIRising anthropogenic pressures in the 20th century have caused extensive habitat loss and fragmentation, threatening biodiversity across ecosystems worldwide. In the boreal forest of Fennoscandia, clear-cut forestry has been a major driver of these changes, resulting in a fragmented landscape of even-aged forest stands. Several studies in recent years have investigated the effect of habitat fragmentation and loss on fungal communities associated with deadwood, but these have mainly focused on visible sporocarps and mushrooms. The effects of forestry on the whole fungal community within deadwood have not been explored as extensively, and especially not in the context of long-term effects. C_LIO_LIWe investigated the effects of clear-cutting on deadwood-inhabiting fungi in boreal Picea abies forests in Norway using ITS2 metabarcoding of sawdust samples collected from 459 logs distributed across 24 paired near-natural and previously clear-cut plots harvested 50 to 90 years ago. C_LIO_LIWhile the overall fungal richness associated with deadwood was similar between the two management types, community composition differed markedly. Plot-scale fungal diversity was linked to deadwood heterogeneity, and composition of the common species was additionally affected by the living tree structure. Nearly all red-listed species were exclusively found in the near-natural forest plots. These findings demonstrate that clear-cut forestry shifts fungal community structure rather than reducing the total number of species and that rare and common species are structured by different environmental drivers but respond in similar ways to large scale disturbance. C_LIO_LISynthesis: Our results highlight the importance of maintaining structural complexity of deadwood in boreal forests for fungal conservation. We found a consistent relationship between deadwood volumes and heterogeneity, and community diversity and species richness. This relationship was consistent even for previously clear-cut forests, showing that retaining structural heterogeneity in managed ecosystems has a positive impact on species diversity. C_LI

ecology↗

Yeasts prefer daycares and molds prefer private homes

Indoor fungi can cause negative health effects due to the production of toxins or volatiles that trigger the immune system of the occupants. To what degree indoor fungi (mycobiomes) differ between buildings with different usage is poorly known. Here, we compare the indoor mycobiomes in 123 childrens daycare centers and 214 private homes throughout Norway, as revealed by metabarcoding of DNA extracted from dust samples collected by community scientists. Although the fungal richness per se was similar in dust samples from daycares and homes, the fungal community composition differed. Yeast fungi, distributed mainly across the orders Saccharomycetales, Filobasidiales and Tremellales, were proportionally more abundant in the daycares, while filamentous fungi, including spore-producing molds such as Aspergillus, Penicillum and Cladosporium, were relatively more abundant in homes. Number of occupants, which is considerably higher in daycares, correlated significantly with the fungal community shift. We hypothesize that the density of occupants and their age distribution drive the systematic difference of yeasts and filamentous fungi in the two building types. ImportanceWorldwide, people spend most of their time indoors; in their homes, workplaces, schools and daycares. The indoor environment can thus largely influence our health. In this study, we revealed that number of occupants and possibly their age distribution are important explanatory variables for the fungal communities of the indoor environments in homes and daycares. Knowledge of these communities are important for further studies on how fungal diversity affect our health.

microbiology↗

The Ribosomal Operon Database (ROD): A full-length rDNA operon database extracted from genome assemblies

Current rDNA reference sequence databases are tailored towards shorter DNA markers, such as parts of the 16/18S marker or the ITS region. However, due to advances in long-read DNA sequencing technologies, longer stretches of the rDNA operon are increasingly used in environmental sequencing studies to increase the phylogenetic resolution. There is, therefore, a growing need for longer rDNA reference sequences. Here, we present the Ribosomal Operon Database (ROD), which includes eukaryotic full-length rDNA operons fished from publicly available genome assemblies. Full-length operons were detected in 34.1% of the 34,701 examined eukaryotic genome assemblies from NCBI. In most cases (53.1%), more than one operon variant was detected, which can be due to intragenomic operon copy variability, allelic variation in non-haploid genomes, or technical errors from the sequencing and assembly process. The highest copy number found was 5,947 in Zea mays. In total, 453,697 unique operons were detected, with 69,480 operon variant clusters remaining after intragenomic clustering at 99% sequence identity. The operon length varied extensively across eukaryotes, ranging from 4,136 to 16,463 bp, which will lead to considerable PCR bias during amplification of the entire operon. Clustering the full-length operons revealed that the different parts (i.e., 18S, 28S, the hypervariable region V4 of 18S, and ITS) provide divergent taxonomic resolution, with 18S and the V4 region being the most conserved. The Ribosomal Operon Database (ROD) will be updated regularly to provide an increasing number of full-length rDNA operons to the scientific community.

genomics↗

Reticulate evolution and rapid development of reproductive barriers upon secondary contact pose challenges for species delineation in a forest fungus

Reproductive barriers within morphospecies of the mushroom-forming fungi tend to be stronger between sympatric lineages, leading to speculation on whether they are being reinforced by selection against hybrids. In this study we use population genomic analyses together with in vitro crosses of a global sample of the wood decay fungus Trichaptum abietinum to investigate reproductive barriers within this morphotaxon and the processes that have shaped them. Our phylogeographic analyses show that the morphospecies is delimited into six major lineages, one in Asia, two in Europe, and three in North America. The two lineages present in Europe are interfertile and admixed, whereas our crosses show that the North American lineages are reproductively isolated. In Asia a more complex pattern appears, with partial intersterility between multiple sub-lineages that likely originated independently and more recently than the reproductive barriers in North America. We found pre-mating barriers in T. abietinum to be moderately correlated with genomic divergence, whereas fitness of the mated hybrids showed a strong positive correlation with increasing genomic divergence. Genome wide association analyses identified candidate genes with programmed cell death annotations, which are known to be involved in intersterility in distantly related fungi. Our demographic modelling and phylogenetic network analyses suggest that reproductive barriers in Trichaptum abietinum could have been reinforced upon secondary contact between lineages that diverged in allopatry during the Pleistocene glacial cycles. Our combination of experimental and genomic approaches demonstrate how T. abietinum is a tractable system for studying speciation mechanims.

evolutionary biology↗

Introgression between highly divergent fungal sister species

To understand how species evolve and adapt to changing environments, it is important to study gene flow and introgression due to their influence on speciation and radiation events. Here, we apply a novel experimental system for investigating these mechanisms using natural populations. The system is based on two fungal sister species with morphological and ecological similarities occurring in overlapping habitats. We examined introgression between these species by conducting whole genome sequencing of individuals from populations in North America and Europe. We assessed genome wide nucleotide divergence and performed crossing experiments to study reproductive barriers. We further used ABBA-BABA statistics together with a network analysis to investigate introgression, and conducted demographic modelling to gain insight into divergence times and introgression events. The results revealed that the species are highly divergent and incompatible in vitro. Despite this, small regions of introgression were scattered throughout the genomes and one introgression event likely involves a ghost population (extant or extinct). This study demonstrates that introgression can be found among divergent species and that population histories can be studied without collections of all the populations involved. Moreover, the experimental system is shown to be a useful tool for research on reproductive isolation in natural populations.

evolutionary biology↗

The indoor mycobiome of daycare centers is affected by occupancy and climate

Many children spend considerable time in daycare centers and may here be influenced by indoor microorganisms, including fungi. In this study, we investigate the indoor mycobiome of 125 daycare centers distributed along strong environmental gradients throughout Norway. Dust samples were collected from doorframes outside and inside buildings using a citizen science sampling approach. Fungal communities in the dust samples were analyzed using DNA metabarcoding of the ITS2 region. We observed a marked difference between the outdoor and indoor mycobiomes. The indoor mycobiome included considerably more yeasts and molds compared to the outdoor samples, with Saccharomyces, Mucor, Malassezia and Penicillium among the most dominant fungal genera. Changes in the indoor fungal richness and composition correlated to numerous variables related to both outdoor and indoor conditions; there was a clear geographic structure in the indoor mycobiome composition that mirrored the outdoor climate, ranging from humid areas in western Norway to drier and colder areas in eastern Norway. Moreover, the number of children in the daycare centers, as well as various building features, influenced the indoor mycobiome composition. We conclude that the indoor mycobiome in Norwegian daycare centers is structured by multiple factors and is dominated by yeasts and molds. This study exemplifies how citizen science sampling enables DNA-based analyses of a high number of samples covering wide geographic areas. ImportanceWith an alarming increase in chronic diseases like childhood asthma and allergies, there is an increased focus on the exposure of young children to indoor biological and chemical air pollutants. Our study of 125 daycares throughout Norway demonstrates that the indoor mycobiome not only reflects co-occurring outdoor fungi but includes a high abundance of yeast and mold fungi with an affinity for indoor environments. A multitude of factors influence the indoor mycobiome in daycares, including building type, inhabitants, as well as the outdoor environment. Many of the detected yeasts and molds are likely associated with the human body, where some have been coupled to allergies and respiratory problems. Our results call for further studies investigating the potential impact of the identified daycare-associated mycobiomes on children health.

microbiology↗

Long-term balancing selection in highly dynamic mating loci generates trans-species polymorphisms in fungi

Balancing selection, an evolutionary force that retains genetic diversity, has been detected in multiple genes and organisms, such as the sexual mating loci in fungi. However, to quantify the strength of balancing selection and define the mating-related genes require a large number of specimens. In tetrapolar basidiomycete fungi, sexual type is determined by two unlinked loci, MATA and MATB. Genes in both loci defines mating type identity, control successful mating and completion of the life cycle. These loci are usually highly diverse. Previous studies have speculated, based on culture crosses, that species of the non-model genus Trichaptum (Hymenochaetales, Basidiomycota) possess a tetrapolar mating system, with multiple alleles. Here, we sequenced a hundred and eighty specimens of three Trichaptum species. We characterized the chromosomal location of MATA and MATB, the molecular structure of MAT regions and their allelic richness. Our sequencing effort was sufficient to molecularly characterize multiple MAT alleles segregating before the speciation event of Trichaptum species. Our analyses suggested that long-term balancing selection has generated trans-species polymorphisms. Mating sequences were classified in different allelic classes based on an amino acid identity (AAI) threshold supported by phylogenetics. The inferred allelic information mirrored the outcome of in vitro crosses, thus allowing us to support the degree of allelic divergence needed for successful mating. Even with the high amount of divergence, key amino acids in functional domains are conserved. The observed allelic classes could potentially generate 14,560 different mating types. We conclude that the genetic diversity of mating in Trichaptum loci is due to long-term balancing selection, with limited recombination and duplication activity. Our large number of sequenced specimens highlighted the importance of sequencing multiple individuals from different species to detect the mating-related genes, the mechanisms generating diversity and the evolutionary forces maintaining them. Author summaryFungi have complex mating systems, and basidiomycete fungi can encode thousands of mating types. Individuals with the same mating type cannot mate. This sexual system has evolved to facilitate sexual mating, increasing the chances to recombine into advantageous allelic combination and prune deleterious alleles. We explored the genomes of hundred and eighty specimens, combined with experimental mating studies of selected specimens, from a non-model organism (Trichaptum). We characterized the genomic regions controlling sex. The mating ability of the specimens confirmed the role of the mating alleles observed in the genomic data. The detailed analyses of many specimens allowed us to observe gene duplication and rearrangements within the mating loci, increasing the diversity within these loci. We supported previous suggestions of balancing selection in this region, an evolutionary force that maintains genomic diversity. These results supports that our fungal specimens are prone to outcross, which might facilitate the adaptation to new conditions.

genomics↗

Wood modification by furfuryl alcohol caused delayed decomposition response in Rhodonia (Postia) placenta

The aim of this study was to investigate differential expression profiles of the brown rot fungus Rhodonia placenta (previously Postia placenta) harvested at several time points when grown on Pinus radiata (radiata pine) and P. radiata with three different levels of modification by furfuryl alcohol, an environmentally benign commercial wood protection system. For the first time the entire gene expression pattern of a decay fungus is followed in untreated and modified wood from initial to advanced stages of decay. Results support the current model of a two-step decay mechanism, with an initial oxidative depolymerization followed by hydrolysis of cell-wall polysaccharides. The wood decay process is finished, and the fungus goes into starvation mode after five weeks when grown on unmodified P. radiata wood. The pattern of repression of oxidative processes and oxalate synthesis found in P. radiata at later stages of decay is not mirrored for the high furfurylation treatment. The high treatment level provided a more unpredictable expression pattern throughout the entire incubation period. Furfurylation does not seem to directly influence the expression of core plant cell wall hydrolyzing enzymes, as a delayed and prolonged, but similar pattern was observed in the P. radiata and the modified experiments. This indicates that the fungus starts a common decay process in the modified wood, but proceeds at a slower pace as access to the plant cell wall polysaccharides is restricted. This is further supported by the downregulation of hydrolytic enzymes for the high treatment level at the last harvest point (mass loss 14%). Moreover, the mass loss does not increase the last weeks. Collectively, this indicates a potential threshold for lower mass loss for highly modified wood.\n\nIMPORTANCEFungi are important decomposers of woody biomass in natural habitats. Investigation of the mechanisms employed by decay fungi in their attempt to degrade wood is important for both the basic scientific understanding of ecology and carbon cycling in nature, and for applied uses of woody materials. For wooden building materials long service life and carbon storage is essential, but decay fungi are responsible for massive losses of wood in service. Thus, optimizing durable wood products for the future are of major importance. In this study we have investigated the fungal genetic response to furfurylated wood, a commercial environmentally benign wood modification approach, that improves service life of wood in outdoor applications. Our results show that there is a delayed wood decay by the fungus as a response to furfurylated wood and new knowledge about the mechanisms behind the delay is provided.

genomics↗