Bacterial rarity in a subarctic stream network: Biodiversity patterns, assembly mechanisms and types of rarity.

Dendritic stream networks are an intriguing subject for exploring the spatial and temporal variability of the rare and common bacterial biosphere, yet very few such studies have been conducted. We sampled riverine bacterioplankton at 13 sites in a subarctic riverine network across 3 years, with five sampling times each year. Ordinations showed a consistent pattern of downstream shift for both rare and abundant subcommunities. We also detected a temporal signal, with seasonal community shifts reflecting changes in water temperature and groundwater contribution, and an inter-annual pattern where the year 2018 differed from other years. Phylogenetic turnover of the rare subcommunity indicated homogeneous selection, whereas the abundant subcommunity was mainly stochastically structured. Transiently rare taxa were the dominant type of rarity with the highest proportion at the headwater regions. The bacterioplankton community was characterized by a small group of core taxa that occurred at most sites with little temporal variation, a very large number of permanently or transiently rare taxa, and taxa shifting through time between the rare and abundant biosphere. While this basic structure could have been detected with less extensive temporal replication, a comprehensive understanding of the rare biosphere in riverine bacterioplankton can only be achieved via inter-annual, spatially replicated sampling that covers the whole stream network.

Weather in two climatic regions shapes the diversity and drives the structure of fungal endophytic community of bilberry (Vaccinium myrtillus L.) fruit.

BACKGROUND: Bilberry (Vaccinium myrtillus L.) is one of the most important economic and natural resources in Northern Europe. Despite its importance, the endophytic fungal community of the fruits has rarely been investigated. Biogeographic patterns and determinants of the fungal diversity in the bilberry fruit are poorly understood, albeit fungal endophytes can have a close relationship with the host plants. Here, we investigated the effect of climatic regions, and their weather conditions within growth season and soil properties on fungal endophytic communities of bilberry fruits collected from northern and southern regions of Finland using high-throughput sequencing technology targeting the internal transcribed spacer 2 ribosomal DNA region for fungi. RESULTS: Species richness and beta diversity (variation in community structure) were higher in the southern compared to the studied northern region. The weather condition of the growth season drove both fungal richness and community structure. Furthermore, abundance of the genera Venturia, Cladosporium, and Podosphaera was influenced by the weather, being different between the south and north regions. CONCLUSIONS: We conclude that diversity and assembly structure of the fungal endophytes in bilberry fruits follow similar patterns as for foliar fungal endophytes, being shaped by various environmental factors, such as the climate and surrounding vegetation.

Growth of floating hook-moss (Warnstorfia fluitans) differs with nutrient and water flow adjustments in greenhouse and cold room conditions.

Floating hook-moss (Warnstorfia fluitans) is a bryophyte growing in northern aquatic and peatland ecosystems. W. fluitans uptakes metals and excessive amounts of nitrogen from wastewater, which suggests that it may have commercial potential for use in phytoremediation. Optimization of growth conditions would allow artificial cultivation of floating hook moss in large quantities for phytoremediation applications. We tested how application of combined nutrient (NPK 7-2-2 ranging from 0.1 to 1 ml per liter of water) and water flow (ranging from 0.15 to 1.9 ml/min) treatments affect growth of W. fluitans in greenhouse conditions. At the end of the experiment, all treatment combinations were subjected to an additional cold room condition at low temperature (0-2 °C) without constant water flow. The moss generally produced biomass in the various treatment combinations. However, contrary to our expectations, we found that increase of nutrients and water flow had a negative effect on the growth of W. fluitans. The highest growth rates in the experiment were detected in the control unit that had no nutrient addition or applied water flow. Our results suggest that cold temperatures are beneficial for W. fluitans growth. Our results show that the commercial production of W. fluitans may not require nutrient or water flow manipulation, at least in the tested scale. Instead, the growth conditions should mimic the natural cold climate conditions of W. fluitans habitats in northern peatlands and/or spring ecosystems.

Bacterial communities in a subarctic stream network: Spatial and seasonal patterns of benthic biofilm and bacterioplankton.

Water-column bacterial communities are assembled by different mechanisms at different stream network positions, with headwater communities being controlled by mass effects (advection of bacteria from terrestrial soils) while downstream communities are mainly driven by environmental sorting. Conversely, benthic biofilms are colonized largely by the same set of taxa across the entire network. However, direct comparisons of biofilm and bacterioplankton communities along whole stream networks are rare. We used 16S rRNA gene amplicon sequencing to explore the spatiotemporal variability of benthic biofilm (2 weeks old vs. mature biofilm) and water-column communities at different network positions of a subarctic stream from early summer to late autumn. Amplicon sequence variant (ASV) richness of mature biofilm was about 2.5 times higher than that of early biofilm, yet the pattern of seasonality was the same, with the highest richness in midsummer. Biofilm bacterial richness was unrelated to network position whereas bacterioplankton diversity was negatively related to water residence time and distance from the source. This pattern of decreasing diversity along the network was strongest around midsummer and diminished greatly as water level increased towards autumn. Biofilm communities were phylogenetically clustered at all network positions while bacterioplankton assemblages were phylogenetically clustered only at the most downstream site. Both early and mature biofilm communities already differed significantly between upstream (1st order) and midstream (2nd order) sections. Network position was also related to variation in bacterioplankton communities, with upstream sites harbouring substantially more unique taxa (44% of all upstream taxa) than midstream (20%) or downstream (8%) sites. Some of the taxa that were dominant in downstream sections were already present in the upmost headwaters, and even in riparian soils, where they were very rare (relative abundance <0.01%). These patterns in species diversity and taxonomic and phylogenetic community composition of the riverine bacterial metacommunity were particularly strong for water-column communities, whereas both early and mature biofilm exhibited weaker spatial patterns. Our study demonstrated the benefits of studying bacterioplankton and biofilm communities simultaneously to allow testing of ecological hypotheses about biodiversity patterns in freshwater bacteria.

Bacterial communities at a groundwater-surface water ecotone: gradual change or abrupt transition points along a contamination gradient?

Microbial communities contribute greatly to groundwater quality, but the impacts of land-use practices on bacteria in groundwaters and groundwater-dependent ecosystems remain poorly known. With 16S rRNA gene amplicon sequencing, we assessed bacterial community composition at the groundwater-surface water ecotone of boreal springs impacted by urbanization and agriculture, using spring water nitrate-N as a surrogate of contamination. We also measured the rate of a key ecosystem process, organic matter decomposition. We documented a recurrent pattern across all major bacterial phyla where diversity started to decrease at unexpectedly low nitrate-N concentrations (100-300 µg L-1 ). At 400 NO3 - -N µg L-1 , 25 bacterial exact sequence variants showed a negative response, resulting in a distinct threshold in bacterial community composition. Chthonomonas, Acetobacterales and Hyphomicrobium were the most sensitive taxa, while only three taxa (Duganella, Undibacterium and Thermoanaerobaculaceae) were enriched due to increased contamination. Decomposition rate responded unimodally to increasing nitrate-N concentration, with a peak rate at ~400 NO3 - -N µg L-1 , parallelly with a major shift in bacterial community composition. Our results emphasize the utility of bacterial communities in the assessment of groundwater-dependent ecosystems. They also call for a careful reconsideration of threshold nitrate values for defining groundwater ecosystem health and protecting their microbial biodiversity.

Host species shape the community structure of culturable endophytes in fruits of wild berry species (Vaccinium myrtillus L., Empetrum nigrum L. and Vaccinium vitis-idaea L.).

Wild berries are interesting research subjects due to their rich sources of health-beneficial phenolic compounds. However, the internal microbial communities, endophytes, associated with the wild berry fruits are currently unknown. Endophytes are bacteria or fungi inhabiting inside plant tissues, and their functions vary depending on the host species and environmental parameters. The present study aimed to examine community composition of fungal and bacterial endophytes in fruits of three wild berry species (bilberry Vaccinium myrtillus L., lingonberry Vaccinium vitis-idaea L. and crowberry Empetrum nigrum L.) and the effects of host plant species and their growth sites on shaping the endophytic communities. We found that the endophytic community structures differed between the berry species, and fungi were predominant over bacteria in the total endophytic taxa. We identified previously unknown endophytic fungal taxa including Angustimassarina, Dothidea, Fellozyma, Pseudohyphozyma, Hannaella coprosmae and Oberwinklerozyma straminea. A role of soluble phenolic compounds, the intracellular components in wild berry fruits, in shaping the endophytic communities is proposed. Overall, our study demonstrates that each berry species harbors a unique endophytic community of microbes.

Habitat patchiness, ecological connectivity and the uneven recovery of boreal stream ecosystems from an experimental drought.

Ongoing climate change is increasing the occurrence and intensity of drought episodes worldwide, including in boreal regions not previously regarded as drought prone, and where the impacts of drought remain poorly understood. Ecological connectivity is one factor that might influence community structure and ecosystem functioning post-drought, by facilitating the recovery of sensitive species via dispersal at both local (e.g. a nearby habitat patch) and regional (from other systems within the same region) scales. In an outdoor mesocosm experiment, we investigated how impacts of drought on boreal stream ecosystems are altered by the spatial arrangement of local habitat patches within stream channels, and variation in ecological connectivity with a regional species pool. We measured basal ecosystem processes underlying carbon and nutrient cycling: (a) algal biomass accrual; (b) microbial respiration; and (c) decomposition of organic matter, and sampled communities of aquatic fungi and benthic invertebrates. An 8-day drought event had strong impacts on both community structure and ecosystem functioning, including algal accrual, leaf decomposition and microbial respiration, with many of these impacts persisting even after water levels had been restored for 3.5 weeks. Enhanced connectivity with the regional species pool and increased aggregation of habitat patches also affected multiple response variables, especially those associated with microbes, and in some cases reduced the effects of drought to a small extent. This indicates that spatial processes might play a role in the resilience of communities and ecosystem functioning, given enough time. These effects were however insufficient to facilitate significant recovery in algal growth before seasonal dieback began in autumn. The limited resilience of ecosystem functioning in our experiment suggests that even short-term droughts can have extended consequences for stream ecosystems in the world's vast boreal region, and especially on the ecosystem processes and services mediated by algal biofilms.

Local geology determines responses of stream producers and fungal decomposers to nutrient enrichment: A field experiment.

We examined how short-term (19 days) nutrient enrichment influences stream fungal and diatom communities, and rates of leaf decomposition and algal biomass accrual. We conducted a field experiment using slow-releasing nutrient pellets to increase nitrate (NO3-N) and phosphate (PO4-P) concentrations in a riffle section of six naturally acidic (naturally low pH due to catchment geology) and six circumneutral streams. Nutrient enrichment increased microbial decomposition rate on average by 14%, but the effect was significant only in naturally acidic streams. Nutrient enrichment also decreased richness and increased compositional variability of fungal communities in naturally acidic streams. Algal biomass increased in both stream types, but algal growth was overall very low. Diatom richness increased in response to nutrient addition by, but only in circumneutral streams. Our results suggest that primary producers and decomposers are differentially affected by nutrient enrichment and that their responses to excess nutrients are context dependent, with a potentially stronger response of detrital processes and fungal communities in naturally acidic streams than in less selective environments.

Groundwater contamination and land drainage induce divergent responses in boreal spring ecosystems.

Degradation of freshwater ecosystems has engendered legislative mandates for the protection and management of surface waters while groundwater-dependent ecosystems (GDEs) have received much less attention. This is so despite biodiversity and functioning of GDEs are currently threatened by several anthropogenic stressors, particularly intensified land use and groundwater contamination. We assessed the impacts of land drainage (increased input of dissolved organic carbon, DOC, from peatland drainage) and impaired groundwater chemical quality (NO3--N enrichment from agricultural or urban land use) on biodiversity and ecosystem functioning in 20 southern Finnish cold-water springs using several taxonomic and functional measures. Groundwater contamination decreased macroinvertebrate and bacterial diversity and altered their community composition. Changes in macroinvertebrate and bacterial communities along the gradient of water-quality impairment were caused by the replacement of native with new taxa rather than by mere disappearance of some of the original taxa. Also species richness of habitat specialist (but not headwater generalist) bryophytes decreased due to impaired groundwater quality. Periphyton accrual rate showed a subsidy-stress response to elevated nitrate concentrations, with peak values at around 2500 µg L-1, while drainage-induced spring water brownification (increased DOC) reduced both periphyton accrual and leaf decomposition rates already at very low concentrations. Our results highlight the underutilized potential of ecosystem-level functional measures in GDE bioassessment as they seem to respond to the first signs of spring ecosystem impairment, at least for the anthropogenic stressors studied by us.