Agostina V. Marano

Zoosporic true fungi in marine ecosystems: a review

Although many species of zoosporic true fungi have been frequently observed and studied in freshwater and soil ecosystems, only three species have been properly identified and partially characterised from brackish and marine ecosystems, namely Rhizophydium littoreum Amon, Thalassochytrium gracilariopsis Nyvall, Pedersen et Longcore and Chytridium polysiphoniae Cohn. These species are either facultative or obligate parasites of marine macroalgae and invertebrates. Also, some species of Olpidium and Rhizophydium are parasites of small marine green algae and diatoms. Although the physiological effects of these pathogens on the growth and metabolism of their hosts are poorly understood, parasitism by C. polysiphoniae possibly affects the rates of photosynthesis and patterns of growth in infected communities of brown algae. Saprobic ecotypes of R. littoreum can also colonise dead-plant and animal substrates. Zoospores from zoosporic true fungi and other groups of microbes possibly provide important food resources for grazing and filter-feeding zooplankton and metazoans in marine ecosystems where the prevalence of disease is high or where accumulated detritus enhances biodiversity in food webs. However, quantitative studies have not yet been attempted. Recently, environmental sampling with molecular techniques has revealed unknown clades of zoosporic true fungi in extreme marine ecosystems. These fungi have been grossly under-sampled and under-studied in marine environments.

Diversity, role in decomposition, and succession of zoosporic fungi and straminipiles on submerged decaying leaves in a woodland stream

Leaf litter is a very important primary source of energy in woodland streams. Decomposition of leaf litter is a process mediated by many groups of microorganisms which release extracellular enzymes for the degradation of complex macromolecules. In this process, true fungi and straminipiles are considered to be among the most active groups, more active than the bacteria, at least during the early stages of the process. Colonization increases the quality of the leaves as a food resource for detritivores. In this way, matter and energy enter detritus-based food chains. Previously, aquatic hyphomycetes were considered to be the major fungal group responsible for leaf litter decomposition. Although zoosporic fungi and straminipiles are known to colonize and decompose plant tissues in various environments, there is scant information on their roles in leaf decomposition. This study focuses on the communities of zoosporic fungi and straminipiles in a stream which are involved in the decomposition of leaves of two plant species, Ligustrum lucidum and Pouteria salicifolia, in the presence of other groups of fungi. A characteristic community dominated by Nowakowskiella elegans, Phytophthora sp., and Pythium sp. was found. Changes in the fungal community structure over time (succession) was observed: terrestrial mitosporic fungi appeared during the early stages, zoosporic fungi, straminipiles, and aquatic Hyphomycetes in early-to-intermediate stages, while representatives of the phylum Zygomycota were found at early and latest stages of the decomposition. These observations highlight the importance of zoosporic fungi and straminipiles in aquatic ecosystems.

Quantitative methods for the analysis of zoosporic fungi

Quantitative estimations of zoosporic fungi in the environment have historically received little attention, primarily due to methodological challenges and their complex life cycles. Conventional methods for quantitative analysis of zoosporic fungi to date have mainly relied on direct observation and baiting techniques, with subsequent fungal identification in the laboratory using morphological characteristics. Although these methods are still fundamentally useful, there has been an increasing preference for quantitative microscopic methods based on staining with fluorescent dyes, as well as the use of hybridization probes. More recently however PCR based methods for profiling and quantification (semi- and absolute) have proven to be rapid and accurate diagnostic tools for assessing zoosporic fungal assemblages in environmental samples. Further application of next generation sequencing technologies will however not only advance our quantitative understanding of zoosporic fungal ecology, but also their function through the analysis of their genomes and gene expression as resources and databases expand in the future. Nevertheless, it is still necessary to complement these molecular-based approaches with cultivation-based methods in order to gain a fuller quantitative understanding of the ecological and physiological roles of zoosporic fungi.

Can zoosporic true fungi grow or survive in extreme or stressful environments

Zoosporic true fungi are thought to be ubiquitous in many ecosystems, especially in cool, moist soils and freshwater habitats which are rich in organic matter. However, some of the habitats where these fungi are found may periodically experience extreme conditions, such as soils in extremely dry, hot and cold climates, acidic and alkaline soils, polluted rivers, anaerobic soil and water, saline soil and water, periglacial soils, oligotrophic soils, tree canopies and hydrothermal vents. It is clear that many ecotypes of zoosporic true fungi have indeed adapted to extreme or stressful environmental conditions. This conclusion is supported by studies in both the field and in the laboratory. Therefore, in our opinion, at least some true zoosporic fungi can be considered to be extremophiles.

Zoosporic parasites infecting marine diatoms - A black box that needs to be opened

Living organisms in aquatic ecosystems are almost constantly confronted by pathogens. Nevertheless, very little is known about diseases of marine diatoms, the main primary producers of the oceans. Only a few examples of marine diatoms infected by zoosporic parasites are published, yet these studies suggest that diseases may have significant impacts on the ecology of individual diatom hosts and the composition of communities at both the producer and consumer trophic levels of food webs. Here we summarize available ecological and morphological data on chytrids, aphelids, stramenopiles (including oomycetes, labyrinthuloids, and hyphochytrids), parasitic dinoflagellates, cercozoans and phytomyxids, all of which are known zoosporic parasites of marine diatoms. Difficulties in identification of host and pathogen species and possible effects of environmental parameters on the prevalence of zoosporic parasites are discussed. Based on published data, we conclude that zoosporic parasites are much more abundant in marine ecosystems than the available literature reports, and that, at present, both the diversity and the prevalence of such pathogens are underestimated.

Frequency, abundance and distribution of zoosporic organisms from Las Cañas stream (Buenos Aires, Argentina).

Zoosporic organisms are common inhabitants of aquatic environments; however there are few ecological studies made for Argentinean streams. In this contribution the taxonomic composition of zoosporic organisms from a stream and their abundance, frequency and diversity on cellulosic baits were analyzed. Samples of water and floating organic matter (vegetable debris) were taken at four dates and different environmental variables (temperature, dissolved oxygen and nutrient concentrations) were measured. Twenty-one taxa were recovered with the baiting technique. Physicochemical fluctuations affected the structure of the studied community; in spring the greatest species richness was related to high nutrient levels whereas in winter the greatest abundance and diversity was related to low water temperature, nutrient levels and well oxygenated conditions.

Ecological functions of zoosporic hyperparasites

Zoosporic parasites have received increased attention during the last years, but it is still largely unnoted that these parasites can themselves be infected by hyperparasites. Some members of the Chytridiomycota, Blastocladiomycota, Cryptomycota, Hyphochytriomycota, Labyrinthulomycota, Oomycota, and Phytomyxea are hyperparasites of zoosporic hosts. Because of sometimes complex tripartite interactions between hyperparasite, their parasite-host, and the primary host, hyperparasites can be difficult to detect and monitor. Some of these hyperparasites use similar mechanisms as their parasite-hosts to find and infect their target and to access food resources. The life cycle of zoosporic hyperparasites is usually shorter than the life cycle of their hosts, so hyperparasites may accelerate the turnaround times of nutrients within the ecosystem. Hyperparasites may increase the complexity of food webs and play significant roles in regulating population sizes and population dynamics of their hosts. We suggest that hyperparasites lengthen food chains but can also play a role in conducting or suppressing diseases of animals, plants, or algae. Hyperparasites can significantly impact ecosystems in various ways, therefore it is important to increase our understanding about these cryptic and diverse organisms.

9 Ecological and Economical Importance of Parasitic Zoosporic True Fungi

Species of zoosporic true fungi have been observed by light microscopy in terrestrial, freshwater, and marine habitats. Most of the described species are saprotrophs or mutualists, but some species are parasites of higher plants, animals, and phytoplankton. Some species play significant ecological roles or cause economically important diseases. A few may cause emerging infectious diseases. Many other species are known only from rDNA sequences obtained from environmental surveys. Unfortunately, in general zoosporic true fungi have been poorly sampled and poorly studied. We predict that many more species will be discovered in the future. Ten examples from three taxonomic groups (Chytridiomycota, Blastocladiomycota, and the Olpidium clade) are discussed in this chapter. The pathosystem model provides an excellent basis for understanding host–parasite relationships and the effect of environmental parameters on these microorganisms. The important ecological roles of zoosporic parasites in food-web dynamics are highlighted. These fungi are particularly well-adapted to their environments because of motile zoospores and resistant sporangia.

Three dimensional quantification of biological samples using micro-computer aided tomography (microCT).

MicroCT is increasingly being used to observe soft animal and plant tissues. Conventional electron and light microscope staining protocols used to enhance the contrast of soft tissues have the potential to be adapted for use in microCT. This would increase the versatility of the microCT beyond improving qualitative observations to facilitating quantitative analysis of soft tissues. This paper describes the development of a culture system and staining protocol which has successfully been used to obtain three dimensional (3-D) quantitative data of filamentous and zoosporic soil fungi. The fungi were grown in an artificial matrix that was developed to simulate the particulate nature of soil. The combination of high contrast staining protocol and use of an X-ray translucent matrix allowed for 3-D qualitative and quantitative analysis of fungal growth. A salient point raised by this study is that the effectiveness of a protocol is reliant on the tissue or cell culture system which includes the composition of the sample, the sampling vessel, the depth of a sample and the combination of stains used. The potential use of this method extends to other fields where distribution and growth patterns in 3-D need to be quantified.

The effects of antifungal substances on some zoosporic fungi (Kingdom Fungi)

Zoosporic fungi constitute a large group of true fungi which inhabit freshwater, brackish, marine and soil ecosystems. In general, very little is known about the effects of antifungal substances on the growth and survival of most species. This review focuses on experimental research with those isolates which have been studied, especially in some species of Synchytrium, Olpidium, Batrachochytrium, Allomyces, Blastocladiella, Neocallimastix. These genera represent genetically diverse groups. Although the research discussed here is restricted to a small sample, some general conclusions can be reached about zoosporic fungi as a whole. Like many other eukaryotic microorganisms, zoosporic fungi are sensitive to a large number of antibiotics, fungicides, surfactants, bacterial metabolites, metabolic poisons, proteins, heavy metals and other antifungal substances. These include substances commonly released into the environment for the control of plant and animal diseases, for increasing production of domestic animals and in the form of waste products from industry. It is possible that the release of antifungal substances into the environment might cause significant changes in the community structure of zoosporic fungi as well as of other groups of microorganisms which play significant roles in food web dynamics and ecosystem complexity. However, this needs documentation by quantitative studies. For these reasons, extensive research on the effects of antifungal substances is much needed.