Parasitic Chytrids Research Articles

Aggregation/disaggregation of microalgal-bacterial flocs in high-rate oxidation ponds is a response to biotic/abiotic-induced changes in microbial community structure

During wastewater treatment by integrated algal pond systems (IAPS), microalgal-bacterial flocs (MaB-flocs) form naturally but periodically disaggregate, resulting in poor settling, low biomass recovery, and reduced effluent quality. This study investigates biotic/abiotic-induced changes in microbial community structure in high-rate algal oxidation ponds (HRAOP) of an IAPS on MaB-floc formation and stability during sewage treatment. Results show that dominance by Pseudopediastrum, Desmodesmus and Micractinium species in spring and summer and the chytrids, Paraphysoderma sp. in spring and Sanchytrium sp. in summer, occurred coincident with enhanced MaB-floc formation and biomass recovery (≥90%). In winter, poor floc formation and low biomass recovery were associated with dominance by Desmodesmus, Chlorella, and the Chlorella-like genus Micractinium. A principal components analysis (PCA) confirmed that combinations of colonial microalgae and associated parasitic chytrids underpin MaB-floc formation and stability in spring and summer and that unicells dominated in winter. Dominance by Thiothrix sp. coincided with floc disaggregation. Thus, changes in season, composition and abundance of colonial microalgae and associated parasitic fungi appeared to impact MaB-floc formation, whereas species composition of the bacterial population and emergence of Thiothrix coincided with floc instability and disaggregation.

Widely used herbicide metolachlor can promote harmful bloom formation by stimulating cyanobacterial growth and driving detrimental effects on their chytrid parasites

Metolachlor (MET) is a widely used herbicide that can adversely affect phytoplanktonic non-target organisms, such as cyanobacteria. Chytrids are zoosporic fungi ubiquitous in aquatic environments that parasitize cyanobacteria and can keep their proliferation in check. However, the influence of organic pollutants on the interaction between species, including parasitism, and the associated ecological processes remain poorly understood. Using the host-parasite system consisting of the toxigenic cyanobacterium Planktothrix agardhii and its chytrid parasite Rhizophydium megarrhizum, we investigated the effects of environmentally relevant concentrations of MET on host-parasite interactions under i) continuous exposure of chytrids and cyanobacteria, and ii) pre-exposure of chytrids. During a continuous exposure, the infection prevalence and intensity were not affected, but chytrid reproductive structures were smaller at the highest tested MET concentration. In the parasite's absence, MET promoted cyanobacteria growth possibly due to a hormesis effect. In the pre-exposure assay, MET caused multi- and transgenerational detrimental effects on parasite fitness. Chytrids pre-exposed to MET showed reduced infectivity, intensity, and prevalence of the infection, and their sporangia size was reduced. Thus, pre-exposure of the parasite to MET resulted in a delayed decline of the cyanobacterial cultures upon infection. After several parasite generations without MET exposure, the parasite recovered its initial fitness, indicating that detrimental effects are transient. This study demonstrates that widely used herbicides, such as MET, could favor cyanobacterial bloom formation both directly, by promoting cyanobacteria growth, and indirectly, by inhibiting their chytrid parasites, which are known to play a key role as top-down regulators of cyanobacteria. In addition, we evidence the relevance of addressing multi-organism systems, such as host-parasite interactions, in toxicity assays. This approach offers a more comprehensive understanding of the effects of pollutants on aquatic ecosystems.

A review of the taxonomic diversity, host-parasite interactions, and experimental research on chytrids that parasitize diatoms.

Diatoms (Bacillariophyta) are a major source of primary production on Earth, generating between 1/4 to 1/2 of all oxygen. They are found in almost all bodies of water, the ice of mountains, the arctic and the antarctic, and soils. Diatoms are also a major source of food in aquatic systems, a key component of the silica cycle, and are carbon capturers in oceans. Recently, diatoms have been examined as sources of biofuels, food, and other economic boons. Chytrids are members of the Kingdom fungi comprising, at a minimum, Chytridiomycota, Blastocladiomycota, and Neocallimastigales. Most chytrids are saprobes, plant pathogens, or parasites, and play an important role in aquatic ecosystems. Chytrid parasitism of diatoms has been reported to cause epidemics of over 90% fatality, though most of the information regarding these epidemics is limited to interactions between just a few hosts and parasites. Given the ubiquity of diatoms, their importance in natural and economic systems, and the massive impact epidemics can have on populations, the relative lack of knowledge regarding parasitism by chytrids is alarming. Here we present a list of the firsthand accounts of diatoms reported parasitized by chytrids. The list includes 162 named parasitic chytrid-diatom interactions, with 63 unique chytrid taxa from 11 genera, and 74 unique diatom taxa from 28 genera. Prior to this review, no list of all documented diatom-chytrid interactions existed. We also synthesize the currently known methods of infection, defense, and experiments examining diatoms and chytrids, and we document the great need for work examining both a greater breadth of taxonomic diversity of parasites and hosts, and a greater depth of experiments probing their interactions. This resource is intended to serve as a building block for future researchers studying diatom-parasite interactions and global planktonic communities in both fresh and marine systems.

The succession of epiphytic microalgae conditions fungal community composition: how chytrids respond to blooms of dinoflagellates

This study aims to investigate the temporal dynamics of the epiphytic protist community on macroalgae, during the summer months, with a specific focus on fungi, and the interactions between zoosporic chytrid parasites and the proliferation of the dinoflagellates. We employed a combination of environmental sequencing techniques, incubation of natural samples, isolation of target organisms and laboratory experiments. Metabarcoding sequencing revealed changes in the dominant members of the epiphytic fungal community. Initially, fungi comprised < 1% of the protist community, mostly accounted for by Basidiomycota and Ascomycota, but with the emergence of Chytridiomycota during the mature phase of the biofilm, the fungal contribution increased to almost 30%. Chytridiomycota became dominant in parallel with an increase in the relative abundance of dinoflagellates in the community. Microscopy observations showed a general presence of chytrids following the peak proliferation of the dinoflagellate Ostreopsis sp., with the parasite, D. arenysensis as the dominant chytrid. The maximum infection prevalence was 2% indicating host-parasite coexistence. To further understand the in-situ prevalence of chytrids, we characterised the dynamics of the host abundance and prevalence of chytrids through co-culture. These laboratory experiments revealed intraspecific variability of D. arenysensis in its interaction with Ostreopsis, exhibiting a range from stable coexistence to the near-extinction of the host population. Moreover, while chytrids preferentially parasitized dinoflagellate cells, one of the strains examined displayed the ability to utilize pollen as a resource to maintain its viability, thus illustrating a facultative parasitic lifestyle. Our findings not only enrich our understanding of the diversity, ecology, and progression of epiphytic microalgal and fungal communities on Mediterranean coastal macroalgae, but they also shed light on the presence of zoosporic parasites in less-explored benthic habitats.

Novel parasitic chytrids infecting snow algae in an alpine snow ecosystem in Japan.

Microbial communities are important components of glacier and snowpack ecosystems that influence biogeochemical cycles and snow/ice melt. Recent environmental DNA surveys have revealed that chytrids dominate the fungal communities in polar and alpine snowpacks. These could be parasitic chytrids that infect snow algae as observed microscopically. However, the diversity and phylogenetic position of parasitic chytrids has not been identified due to difficulties in establishing their culture and subsequent DNA sequencing. In this study, we aimed to identify the phylogenetic positions of chytrids infecting the snow algae, Chloromonas spp., bloomed on snowpacks in Japan. By linking a microscopically picked single fungal sporangium on a snow algal cell to a subsequent sequence of ribosomal marker genes, we identified three novel lineages with distinct morphologies. All the three lineages belonged to Mesochytriales, located within "Snow Clade 1", a novel clade consisting of uncultured chytrids from snow-covered environments worldwide. Additionally, putative resting spores of chytrids attached to snow algal cells were observed. This suggests that chytrids may survive as resting stage in soil after snowmelt. Our study highlights the potential importance of parasitic chytrids that infect snow algal communities.

Infection strategies of different chytrids in a diatom spring bloom

Abstract Diatom blooms are often accompanied by an increase in parasitic chytrids that kill the host cells, which they are infecting, and can contribute to the decline of the bloom. However, host specificity and range of these chytrids are currently poorly understood. Low host specificity would enable the parasites to opportunistically infect any diatom species, while specialisation on infecting specific high‐biomass species could result in high prevalence and rapid spread of infection. We investigated such host–parasite interactions by monitoring the diverse diatom spring bloom in lake Erken using amplicon sequencing. We also performed infection experiments with two different, newly isolated chytrid species and several diatom cultures from the bloom. Chytridiomycota displayed the highest relative abundance of all parasitic lineages and were probably physically attached to larger organisms. Since the chytrids reached maximum abundance shortly after a peak in diatom reads, they were probably infecting these important primary producers. Phylogenetic analyses of the isolated chytrid strains identified them as members of the classes Rhizophydiales and Lobulomycetales. The infection experiments revealed high host specificity in these two chytrids targeting different diatom species. The experimental results supported statistical analyses of the environmental sequencing data, which suggested the presence of two different infection strategies: the most abundant chytrid species were specialised on infecting dominant diatom genera (i.e. Stephanodiscus, Aulacoseira, Asterionella), while rarer chytid species infected a range of less abundant diatoms.

High prevalence of parasitic chytrids infection of glacier algae in cryoconite holes in Alaska

Glacier algae, which are photosynthetic microbes growing on ice, considerably reduce the surface albedo of glaciers and accelerate their melting rate. Although the growth of glacier algae can be suppressed by parasitic chytrids, the impact of chytrids on algal populations is still largely unknown. In this study, we described the morphology of the chytrid infecting the glacier alga Ancylonema nordenskioeldii and quantified the prevalence of infection in different habitats on a mountain glacier in Alaska, USA. Microscopic observations revealed three different morphological types of chytrids with distinct rhizoid shapes. Variations in the size of the sporangia were probably because of their different growth stages, indicating that they actively propagated on the glacier. The prevalence of infection did not vary among sites with different elevations but was substantially higher in cryoconite holes (20%) than on ice surfaces (4%) at all sites. This indicates that cryoconite holes are hot spots for chytrid infections of glacier algae, and the dynamics of cryoconite holes might affect the host–parasite interactions between chytrids and the glacier algae, which may in turn alter surface albedo and ice melting.

Species composition of Cyanoprokaryota in the summer phytoplankton of 55 lakes and reservoirs, sampled in Bulgaria in the years 2018, 2019, 2021 and 2023

The paper presents detailed results on cyanoprokaryote diversity in the summer phytoplankton of 55 lakes, small and large reservoirs sampled in Bulgaria in the frame of three joined projects related to algal blooms threat to human health and national security. The phytoplankton of twenty from these selected waterbodies was sampled for first time. In total, 185 species and one variety from 55 genera have been identified, 54 of which (29%) were novel for the country. The average contribution of cyanophytes to the phytoplankton per site was 8 species (or 44% of the total biodiversity), reaching in some sites 80%. According to the morphology, the recorded algae were distributed as follows: 83 coccal, 52 non-heterocytous filamentous and 51 heterocytous filamentous cyanoprokaryotes. Their average contribution to the phytoplankton diversity was estimated as 4 coccal, 2 non-heterocytous and 2 heterocytous species per site. Most of the cyanoprokaryote species (94, or 51%) were recorded only once, even in the case of wetlands and sites which have been repeatedly visited during all sampling campaigns. These 96 species embrace 39 coccal, 23 non-heteroctous and 32 heterocytous forms. No species was found in all studied waterbodies. The most widely spread species were Microcystis aeruginosa (19 records), Planktolyngbya limnetica (17 records), Aphanizomenon klebahnii (16 records), Microcystis wesenbergii (16 records), Aphanocapsa delicatissima (13 records), Cuspidothrix issatschenkoi (12 records), Coelomoron pusillum (11 records), Pseudanabaena limnetica (11 records), Anagnostidinema amphibium (10 records), Raphidiopsis raciborskii (10 records), Limnococcus limneticus (10 records), and separate cells of Microcystis as well (15 records). The record of the chytrid parasite Rhizosiphon anabaenae on separate trichomes of Sphaerospermopsis aphanizomenoides is described.

Mucilage protects the planktonic desmid Staurodesmus sp. against parasite attack by a chytrid fungus.

Zoosporic fungi of the phylum Chytridiomycota are ubiquitous parasites of phytoplankton in aquatic ecosystems, but little is known about phytoplankton defense strategies against parasitic chytrid attacks. Using a model chytrid-phytoplankton pathosystem, we experimentally tested the hypothesis that the mucilage envelope of a mucilage-forming desmid species provides protection against the parasitic chytrid Staurastromyces oculus. Mucilage-forming Staurodesmus cells were not accessible to the chytrid, whereas physical removal of the mucilage envelope rendered the same Staurodesmus sp. strain equally susceptible to chytrid infections as the original non-mucilage-forming host Staurastrum sp. Epidemic spread of the parasite only occurred in Staurastrum sp., whereas non-mucilage-bearing Staurodesmus sp. allowed for co-existence of host and parasite, and mucilage-bearing Staurodesmus sp. caused parasite extinction. In addition to the mucilage defense barrier, we also demonstrate the ability of both Staurastrum sp. and Staurodesmus sp. to resist infection by preventing chytrid development while still remaining viable and being able to reproduce and thus recover from an infection. This study extends our knowledge on phytoplankton defense traits and the functional role of mucilage in phytoplankton as a physical barrier against fungal parasites.

Moving beyond standard toxicological metrics: The effect of diclofenac on planktonic host-parasite interactions

Pharmaceuticals are increasingly released into surface waters and therefore ubiquitous in aquatic systems. While pharmaceuticals are known to influence species interactions, their effect on host-parasite interactions is still underexplored despite potential ecosystem-level consequences. Here, we ask whether diclofenac, a widely used non-steroid anti-inflammatory drug, affects the interaction between a phytoplankton host (Staurastrum sp.; green alga) and its obligate fungal parasite (Staurastromyces oculus; chytrid fungus). We hypothesized that the effect of increasing diclofenac concentration on the host-parasite system depends on parasite exposure. We assessed acute and chronic effects of a wide range of diclofenac concentrations (0–150 mg/L) on host and parasite performance using a replicated long gradient design in batch cultures. Overall system response summarizing parameters related to all biotic components in an experimental unit i.e., number of bacteria and phytoplankton host cells along with photosynthetic yield (a measure of algal cell fitness), depended on diclofenac concentration and presence/absence of parasite. While host standing biomass decreased at diclofenac concentrations >10 mg/L in non-parasite-exposed treatments, it increased at ≥10 mg/L in parasite-exposed treatments since losses due to infection declined. During acute phase (0–48 h), diclofenac concentrations <0.1 mg/L had no effect on host net-production neither in parasite-exposed nor non-parasite-exposed treatments, but parasite infection ceased at 10 mg/L. During chronic phase (0–216 h), host net-production declined only at concentrations >10 mg/L in non-parasite-exposed cultures, while it was overall close to zero in parasite-exposed cultures. Our results suggest that chytrid parasites are more sensitive to diclofenac than their host, allowing a window of opportunity for growth of phytoplankton hosts, despite exposure to a parasite. Our work provides a first understanding about effects of a pharmaceutical on a host-parasite interaction beyond those defined by standard toxicological metrics.

Complex effects of chytrid parasites on the growth of the cyanobacterium Planktothrix rubescens across interacting temperature and light gradients

Chytrids are important drivers of aquatic ecosystems as phytoplankton parasites. The interaction between these parasites and their hosts are shaped by abiotic factors such as temperature and light. Here, we performed a full-factorial experiment to study how temperature and light interact to affect the dynamics of the bloom-forming toxic cyanobacterium Planktothrix rubescens and its chytrid parasite. We used a dynamic host-parasite model to explore how temperature and light affect long term dynamics. At low temperatures, chytrids do not survive. Higher light and temperature levels stimulated both phytoplankton and chytrid growth, with complex effects on their dynamics. Model exploration indicates that increasing temperature and light shifts equilibrium outcomes from P. rubescens persisting alone to stable coexistence and then to limit cycles. This provides an alternative biological explanation for why P. rubescens is mainly found in the relatively cold and dark lake metalimnion – it may enable avoidance of its parasite. Our study emphasizes the importance of investigating how abiotic factors interact with biotic interactions to drive complex outcomes.

Fate of hepatotoxin microcystin during infection of cyanobacteria by fungal chytrid parasites

Chytrid parasites are increasingly recognized as ubiquitous and potent control agents of phytoplankton, including bloom-forming toxigenic cyanobacteria. In order to explore the fate of the cyanobacterial toxin microcystins (MCs) and assess potential upregulation of their production under parasite attack, a laboratory experiment was conducted to evaluate short- and long-term variation in extracellular and intracellular MC in the cyanobacteria Planktothrix agardhii and P. rubescens, both under chytrid infection and in the presence of lysates of previously infected cyanobacteria. MCs release under parasite infection was limited and not different to uninfected cyanobacteria, with extracellular toxin shares never exceeding 10%, substantially below those caused by mechanical lysis induced by a cold-shock. Intracellular MC contents in P. rubescens under infection were not significantly different from uninfected controls, whereas infected P. agardhii showed a 1.5-fold increase in intracellular MC concentrations, but this was detected within the first 48 hours after parasite inoculation and not later, indicating no substantial MC upregulation in cells being infected. The presence of lysates of previously infected cyanobacteria did not elicit higher intracellular MC contents in exposed cyanobacteria, speaking against a putative upregulation of toxin production induced via quorum sensing in response to parasite attack. These results indicate that chytrid epidemics can constitute a bloom decay mechanism that is not accompanied by massive release of toxins into the medium.

Seasonality of parasitic and saprotrophic zoosporic fungi: linking sequence data to ecological traits

Zoosporic fungi of the phylum Chytridiomycota (chytrids) regularly dominate pelagic fungal communities in freshwater and marine environments. Their lifestyles range from obligate parasites to saprophytes. Yet, linking the scarce available sequence data to specific ecological traits or their host ranges constitutes currently a major challenge. We combined 28 S rRNA gene amplicon sequencing with targeted isolation and sequencing approaches, along with cross-infection assays and analysis of chytrid infection prevalence to obtain new insights into chytrid diversity, ecology, and seasonal dynamics in a temperate lake. Parasitic phytoplankton-chytrid and saprotrophic pollen-chytrid interactions made up the majority of zoosporic fungal reads. We explicitly demonstrate the recurrent dominance of parasitic chytrids during frequent diatom blooms and saprotrophic chytrids during pollen rains. Distinct temporal dynamics of diatom-specific parasitic clades suggest mechanisms of coexistence based on niche differentiation and competitive strategies. The molecular and ecological information on chytrids generated in this study will aid further exploration of their spatial and temporal distribution patterns worldwide. To fully exploit the power of environmental sequencing for studies on chytrid ecology and evolution, we emphasize the need to intensify current isolation efforts of chytrids and integrate taxonomic and autecological data into long-term studies and experiments.

Diversity, activity, and abundance of benthic microbes in the Southeastern Mediterranean Sea.

Benthic microbes are key organisms in the oligotrophic Southeastern Mediterranean Sea (SEMS), yet their abundance, activity, and diversity in this rapidly changing basin are not fully understood. We investigated the prokaryotic and microfungal communities throughout years 2018-2020 at 27 stations (6-1900m water depths, down to 20cm below the sediment surface), in two transects with distinct downslope transport regimes, and along the eutrophic coastline. We estimated microbial abundance with flow cytometry, secondary production as leucine assimilation, and sequenced marker genes (the 16S rRNA and internal transcribed spacer) to assess diversity indices. The highest abundance (0.21 × 108cellsgr-1 sediment) was estimated at slope stations where we assumed substantial transport rates and found an accumulation of organic carbon. Secondary production was the highest nearshore (12±4ngCgr-1 h-1), and markedly declined offshore (0.5±0.9ngCgr-1 h-1). Populations of archaea (dominant Nitrososphaeria and Nanoarchaeia) and diverse bacteria were stable over three years, and taxonomic composition was dictated mainly by depth gradients. Saprotrophic and pathotrophic microfungi Ascomycota (70%±23%)andBasidiomycota (16%±18%)wereprevalent, whereas parasitic chytrids were abundant nearshore. Our results highlight the role of downslope transport, which enriched the typical deep-sea communities with anaerobic lineages, in shaping microbial populations near the continental slope.

Revisiting the isolation source after half a century: Emericellopsis mirabilis on a yellow-green alga.

Fungi-algae interactions, such as lichen-forming fungi and parasitic chytrids on phytoplankton, are common in ecosystems. In contrast, interactions between filamentous fungi and soil algae that can be observed with the naked eye have been given little attention and remain unexplored. Here, we report a fungus that was associated with a visible symptom of dead algae on a soil surface in Sugadaira-kogen, Nagano, central Japan. Acremonium-like conidiophores were growing on vesicles and dead bodies of a yellow-green alga, Botrydium granulatum. The fungus was identified as Emericellopsis mirabilis based on its morphology by microscopic observation, phylogenetic analysis, and the similarity of the isolation substrate with the first description of the species. Co-culture experiments showed a filamentous cell differentiation of the alga by the fungus, but no harmful or beneficial effects on algal growth. Therefore, we speculate that E. mirabilis is a facultative parasite of B. granulatum under natural conditions.

Temporal variation in community structure of zoosporic fungi in Lake Biwa, Japan

Zoosporic fungi play an important role in aquatic environments, but their diversity, especially that of parasitic fungi of phytoplankton, has still not been fully revealed. We conducted monthly analyses of the community structure of zoosporic fungi at a pelagic site in Lake Biwa, Japan, from May to December 2016. Metabarcoding analysis, targeted to a large subunit region of ribosomal DNA in the nano-size fraction of particles (2-20 µm), was carried out on the samples. We also counted large phytoplankton and chytrid sporangia attached to the hosts. We detected 3 zoosporic fungal phyla (Blastocladiomycota, Chytridiomycota and Cryptomycota) within 107 operational taxonomic units (OTUs), in which Chytridiomycota was the most diverse and abundant phylum. Few fungal OTUs overlapped between months, and specific communities were detected in each month. These results showed that diverse zoosporic fungi with high temporal variability inhabited the lake. Five large phytoplankton species were found to be infected by chytrids: Staurastrum dorsidentiferum, S. rotula, Closterium aciculare, Asterionella formosa and Aulacoseira granulata. Some chytrids were detected by metabarcoding analysis: Zygophlyctis asterionellae infecting A. formosa, Staurastromyces oculus infecting S. dorsidentiferum and Pendulichytrium sphaericum infecting A. granulata. One OTU detected in association with infected C. aciculare by microscopic counting might have been an obligate parasitic chytrid of C. aciculare. The results indicated that a combination of metabarcoding and microscopic analysis revealed more information on zoosporic fungi, including those that are parasitic.

Parasitoid chytridiomycete Ericiomyces syringoforeus gen. et sp. nov. has unique cellular structures to infect the host

Many fungi have been identified as pathogens of marine algae. Among them, Chytridiomycota have been revealed as relatively highly abundant, but much of the diversity known within these groups is almost entirely based on environmental sequencing data. Here, we present a novel chytridiomycete genus and species, characterized by light microscopical observations, ultrastructure, and molecular phylogenetic analysis of the parasitic chytrid of brackish-water dinoflagellate Kryptoperidinium foliaceum from the Baltic Sea. Phylogenetic analysis of rDNA sequences and the ultrastructure of the strain reveals that it represents a new family in the order Rhizophydiales. Ericiomyces syringoforeus gen. et sp. nov. is a parasitoid with a life cycle composed by zoospores, which attach to the host, encyst, and produce a rhizoidal system (haustorium). Unlike typical Rhizophydiales chytrids, sporangium develops as a lateral outgrowth of the encysted zoospore. The ultrastructural study revealed at least two unique traits: the syringe-like organelle in the cyst, which supposed to paralyze the host, and funnel-shaped structure anchoring sporangium in the host wall. Sporangium matures and produces new zoospores within 3 days. Multiple infections are common and then the life cycle is 1–2 days shorter compared to the duration when a single infection occurred. Cross-infection experiments showed that E. syringoforeus could only infect dinoflagellates, being K. foliaceum highly susceptible to infection by the chytrid parasitoid. The effects of some fungal epidemics on populations of Kryptoperidinium are discussed.

Single dominant diatom can host diverse parasitic fungi with different degree of host specificity

AbstractRecent molecular surveys revealed an unexpected diversity of mycoplankton in lakes and oceans. The early diverging fungal lineages are known to be prominent parasites of phytoplankton. However, due to missing fungal reference data, their identity and ecology remain mostly unknown. To overcome this problem, we combined single‐spore‐based DNA barcoding during one sampling season of the eutrophic Lake Inba (Japan), which is dominated by two diatoms. By linking microscopically picked single fungal spore on two diatoms to subsequent DNA barcoding of ribosomal maker genes, we identified 12 distinct lineages, affiliated not only to the known parasitic phylum Chytridiomycota (chytrids), but also to the enigmatic phyla Rozellomycota and Aphelidiomycota. The detected Rozellomycota could be a hyperparasite of parasitic chytrid infecting diatoms. Host specificity appeared to be different among clades. However, the barcoding of single‐spore DNA could not clearly prove host specificity due to the limited number of samples and resolution in the targeted gene regions. The degree of host specificity was thus confirmed by the cross‐infection experiments. Five chytrid strains were generalists infecting all four diatoms strains, while the remaining three strains of chytrids were specialists exhibiting host preferences. Additional growth experiments indicated a trade‐off between host specificity and growth rate, so that generalists with more possible hosts grew slower than specialists with one host. Our results suggest that the combination of microscopy with single‐spore‐based barcoding is a promising approach to evaluate the complex host‐parasite interactions, while infection experiments can verify the interactions and shed light on the underlying ecological principles.

Parasitic Chytrids Upgrade and Convey Primary Produced Carbon During Inedible Algae Proliferation

Microbial parasites have only recently been included in planktonic food web studies, but their functional role in conveying dietary energy still remains to be elucidated. Parasitic fungi (chytrids) infecting phytoplankton may constitute an alternative trophic link and promote organic matter transfer through the production of dissemination zoospores. Particularly, during proliferation of inedible or toxic algal species, such as large Cyanobacteria fostered by global warming, parasites can constitute an alternative trophic link providing essential dietary nutrients that support somatic growth and reproduction of consumers. Using phytoplankton-parasites associations grown under laboratory controlled conditions we assessed the fatty acids and biochemical composition of species with different nutritional quality and followed the metabolic pathway from the algal host and their parasites zoospores using compound-specific stable isotope analysis. This study demonstrated that chytrids are trophic upgraders able to retain essential nutrients that can be transferred to upper trophic levels both in terms of organic matter quantity and nutritional quality. Through the production of zoospores, nutritionally important long-chain polyunsaturated fatty acids that can be consequently assimilated by consumers. We conclude that parasitism at the base of aquatic food webs may represent a crucial trophic link for dietary nutrients and essential biomolecules alternative to herbivory or bacterivory, which can be particularly crucial during the proliferation of inedible or nutritionally inadequate algal species fostered by climate change.

Light intensity and spectral distribution affect chytrid infection of cyanobacteria via modulation of host fitness.

Light gradients are an inherent feature in aquatic ecosystems and play a key role in shaping the biology of phytoplankton. Parasitism by chytrid fungi is gaining increasing attention as a major control agent of phytoplankton due to its previously overlooked ubiquity, and profound ecological and evolutionary consequences. Despite this interest, if and how light conditions modulate phytoplankton chytridiomycosis remains poorly studied. We investigated life-history traits of a chytrid parasite, Rhizophydium megarrhizum, under different light intensities and spectral compositions when infecting two closely related planktonic cyanobacteria with different light-harvesting strategies: Planktothrix rubescens and P. agardhii. In general, parasite transmission was highest under light conditions (both intensity and quality) that maximized growth rates for uninfected cyanobacteria. Chytrid encystment on hosts was significantly affected by light intensity and host strain identity. This likely resulted from higher irradiances stimulating the increased discharge of photosynthetic by-products, which drive parasite chemotaxis, and from strain-specific differences at the cell-surface. Comparisons of parasite transmission and host growth rates under different light conditions suggest the potential for epidemic development at higher irradiances, whereas host and parasite could coexist without epidemic outbreaks at lower light levels. These results illustrate the close relationship between parasite transmission and host fitness, which is ultimately modulated by the external environment.