Mycoloop

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The mycoloop is a trophic pathway in aquatic food webs where parasitic fungi, particularly chytrids, facilitate the transfer of nutrients and energy from large, inedible phytoplankton (algae) to zooplankton. This process enhances nutrient cycling and supports higher trophic levels in aquatic ecosystems.

Contents

Chytrids infect large, inedible phytoplankton, such as diatoms or cyanobacteria, and produce zoospores (free-living, motile spores, 2–5 μm in diameter). These zoospores are rich in nutrients like polyunsaturated fatty acids (PUFAs) and cholesterol, making them an excellent food source for zooplankton, such as Daphnia and rotifers. By consuming the zoospores or fragmented phytoplankton, zooplankton gain access to nutrients that would otherwise be unavailable from inedible phytoplankton, creating the trophic link called the mycoloop. In this manner, the mycoloop channels carbon, phosphorus, and other nutrients from phytoplankton to zooplankton, bypassing the limitations of inedible phytoplankton.

The mycoloop can influence phytoplankton blooms by reducing host populations (via parasitism) and supporting zooplankton growth, potentially stabilizing aquatic food webs. It can also influence the carbon cycle by altering carbon fluxes, reducing the sinking of large phytoplankton and redirecting carbon to higher trophic levels.

The concept of the mycoloop was developed by Maiko Kagami et al. in 2007. [1] The term "mycoloop" combines myco (referring to fungi, specifically chytrids) with loop (referring to the cycle of nutrient transfer). The discovery of the mycoloop, and its potential impact on nutrient cycling indicates the importance of fungal-algal interactions in natural systems. Chytrids have also been reported to stabilize food webs, while also reducing the amount of organic material that reaches benthic environments. [2] [3]

Background

Most food web studies do not incorporate what is perhaps the most common trophic interaction - parasitism. [4] Despite their ubiquity, parasites are often overlooked because of their cryptic nature, the difficulties in quantifying their effects, and their assumed low biomass. [5] However, they can account for greater biomass than predators [6] and participate in the majority of trophic links. [7] Parasites can modulate trophic flows in a number of ways. They can drive reductions in host biomass, not only by increasing host mortality rates, but also by influencing growth, fecundity, nutritional status, susceptibility to predation, or behaviour. [8] While their role as consumers is better known, parasites can also be prey for other organisms. They can be consumed together with their host (i.e. concomitant predation) or as free living life stages. Given the enormous reproductive output of parasites, free living infecting stages potentially constitute a significant nutrient source and can account for a substantial transfer of material and energy to higher trophic levels. [9] [10] [11]

Chytrids are a type of microscopic fungi belonging to the phylum Chytridiomycota. These fungi are primarily aquatic or found in moist environments. [12] Chytrids can be saprophytic (decomposing organic matter), parasitic (infecting plants, algae, or animals), or mutualistic, and play key ecological roles in breaking down organic material and nutrient cycling. [13] Notably, the chytrid Batrachochytrium dendrobatidis causes chytridiomycosis, a deadly disease in amphibians, contributing to global population declines. [14] [15] They are unusual among fungi in that they reproduce with motile spores, driven by flagella, called zoospores. [16] [17] Most chytrids do not sexually reproduce. Asexual reproduction occurs through the release of zoospores. [16]

Chytrid zoospores are tiny fungal spores which in the ocean can have densities up to a billion per litre. They are rich in nutrients making them excellent food sources for zooplankton. Chytrid zoospores.jpg
Chytrid zoospores are tiny fungal spores which in the ocean can have densities up to a billion per litre. They are rich in nutrients making them excellent food sources for zooplankton.
Pennate diatom from an Arctic meltpond, infected with two chytrid-like zoosporangium fungal pathogens (in false-colour red). Scale bar = 10 mm. Pennate diatom infected with two chytrid-like fungal pathogens.png
Pennate diatom from an Arctic meltpond, infected with two chytrid-like zoosporangium fungal pathogens (in false-colour red). Scale bar = 10 μm.

Saprotrophic chytrids decompose inedible organic matter releasing zoospores that zooplankton consume, further contributing to nutrient cycling. Zooplankton grazing on zoospores may suppress chytrid outbreaks, regulating parasite populations. The mycoloop can stabilise ecosystem by alleviating competition among phytoplankton and supporting zooplankton production, especially in nutrient-rich environments. [3]

Studies show chytrid zoospores, which can have densities up to a billion spores per litre, are a high-quality food source, doubling zooplankton feeding rates compared to uninfected phytoplankton. The mycoloop is significant both in freshwater lakes and marine environments, with chytrids like Zygorhizidium facilitating nutrient transfer from algae like Asterionella to zooplankton like Daphnia. [20] [3]

Parasitic fungi derive nutrients from living hosts, often causing harm. However, fungi have many other ecological roles they can play apart from being parasitic. For example, they can be mycorrhizal (forming mutualistic relationships with plants), endophytic (living inside plants without causing harm), lichenized (forming symbiotic relationships with algae or cyanobacteria), or saprotrophic (breaking down dead organic matter). Some fungi even switch roles depending on environmental conditions or host availability.

Mycoplankton

Mycoplankton are saprotrophic or parasitic members of the plankton communities of marine and freshwater ecosystems. [21] [22] [23] They are composed of filamentous free-living fungi and yeasts that are associated with planktonic particles or phytoplankton. [24] Similar to bacterioplankton, these aquatic fungi play a significant role in heterotrophic mineralization and nutrient cycling. [25] Mycoplankton can be up to 20 mm in diameter and over 50 mm in length, [26] though mostly they are microscopic. [27] [28] A typical litre of seawater contains between one- and ten-million fungal cells. [29] [23] The number is greater in coastal ecosystems and estuaries due to nutritional runoff from terrestrial communities.

Aquatic fungi are found in a myriad of ecosystems, from mangroves, to wetlands, to the open ocean. [30] The greatest diversity and number of species of mycoplankton is found in surface waters (< 1000 m), and the vertical profile depends on the abundance of phytoplankton. [31] [32] Furthermore, this difference in distribution may vary between seasons due to nutrient availability. [33] Aquatic fungi survive in a constant oxygen deficient environment, and therefore depend on oxygen diffusion by turbulence and oxygen generated by photosynthetic organisms. [34]

Aquatic fungi consist mostly of tiny mycoplankton (microfungi), yeast, or mobile zoospores, that can recycle organic matter through the mycoloop process, which involving parasiting plankton. [35] Instead of directly building biomass, decomposers break organic nutrients down into inorganic forms which can be recycled (an approach which metabolically can be costly). [35]

Mycoloop dynamics

Parasitic chytrids can transfer material from large inedible phytoplankton to zooplankton. Chytrids zoospores are excellent food for zooplankton in terms of size (2–5 μm in diameter), shape, nutritional quality (rich in polyunsaturated fatty acids and cholesterols). Large colonies of host phytoplankton may also be fragmented by chytrid infections and become edible to zooplankton. [36]

Parasitic fungi, as well as saprotrophic fungi, directly assimilate phytoplankton organic carbon. By releasing zoospores, the fungi bridge the trophic linkage to zooplankton, known as the mycoloop. By modifying the particulate and dissolved organic carbon, they can affect bacteria and the microbial loop. These processes may modify marine snow chemical composition and the subsequent functioning of the biological carbon pump. [37] [38]

Diagram of a mycoloop (fungus loop) Parasitic chytrids can transfer material from large inedible phytoplankton to zooplankton. Chytrids zoospores are excellent food for zooplankton in terms of size (2-5 mm in diameter), shape, nutritional quality (rich in polyunsaturated fatty acids and cholesterols). Large colonies of host phytoplankton may also be fragmented by chytrid infections and become edible to zooplankton. Parasitic chytrids as a mycoloop.jpg
Diagram of a mycoloop (fungus loop)
Parasitic chytrids can transfer material from large inedible phytoplankton to zooplankton. Chytrids zoospores are excellent food for zooplankton in terms of size (2–5 μm in diameter), shape, nutritional quality (rich in polyunsaturated fatty acids and cholesterols). Large colonies of host phytoplankton may also be fragmented by chytrid infections and become edible to zooplankton.
Possible other mycoloops in aquatic environments Saprotrophic chytrids may also play important roles in aquatic food webs, by decomposing inedible organic material such as pollens. Zoospores released from pollen may be consumed by zooplankton, functioning as another "mycoloop." In addition to chytrids, other zoosporic fungi or fungal-like protists, such as Cryptomycota and Labyrinthulomycota, can infect phytoplankton or consume large inedible organic material, which may be grazed by zooplankton in freshwater and marine environments. Possible mycoloops in aquatic environments.webp
Possible other mycoloops in aquatic environments
Saprotrophic chytrids may also play important roles in aquatic food webs, by decomposing inedible organic material such as pollens. Zoospores released from pollen may be consumed by zooplankton, functioning as another "mycoloop." In addition to chytrids, other zoosporic fungi or fungal-like protists, such as Cryptomycota and Labyrinthulomycota, can infect phytoplankton or consume large inedible organic material, which may be grazed by zooplankton in freshwater and marine environments.
Mycoloop links between phytoplankton and zooplankton Chytrid-mediated trophic links between phytoplankton and zooplankton (mycoloop). While small phytoplankton species can be grazed upon by zooplankton, large phytoplankton species constitute poorly edible or even inedible prey. Chytrid infections on large phytoplankton can induce changes in palatability, as a result of host aggregation (reduced edibility) or mechanistic fragmentation of cells or filaments (increased palatability). First, chytrid parasites extract and repack nutrients and energy from their hosts in form of readily edible zoospores. Second, infected and fragmented hosts including attached sporangia can also be ingested by grazers (i.e. concomitant predation). Mycoloop links between phytoplankton and zooplankton.jpg
Mycoloop links between phytoplankton and zooplankton
Chytrid‐mediated trophic links between phytoplankton and zooplankton (mycoloop). While small phytoplankton species can be grazed upon by zooplankton, large phytoplankton species constitute poorly edible or even inedible prey. Chytrid infections on large phytoplankton can induce changes in palatability, as a result of host aggregation (reduced edibility) or mechanistic fragmentation of cells or filaments (increased palatability). First, chytrid parasites extract and repack nutrients and energy from their hosts in form of readily edible zoospores. Second, infected and fragmented hosts including attached sporangia can also be ingested by grazers (i.e. concomitant predation).
Mycoloop with diatom and rotifer The food web system includes the inedible diatom (Synedra), the obligate parasitic consumer of the diatom (chytrid) with a sessile (sporangium) and a motile (zoospore) life stage, and the rotifer (Keratella), which can consume the chytrid zoospores but not the host diatom. While Synedra is inedible to Keratella, its nutrients may still be transferred to the rotifer via infection propagules (zoospores). Mycoloop with diatom and rotifer.webp
Mycoloop with diatom and rotifer
The food web system includes the inedible diatom (Synedra), the obligate parasitic consumer of the diatom (chytrid) with a sessile (sporangium) and a motile (zoospore) life stage, and the rotifer (Keratella), which can consume the chytrid zoospores but not the host diatom. While Synedra is inedible to Keratella, its nutrients may still be transferred to the rotifer via infection propagules (zoospores).
The central role played by pelagic fungi, both parasitic and saprotrophic in the mycoloop, and saprotrophic fungi as active contributors to the microbial loop. The activity of heterotrophic microbes, including pelagic fungi, has far-reaching global implications for fisheries (i.e., the amount of carbon that will ultimately flow to higher trophic levels) and climate change (i.e., the amount of carbon that will be sequestered in the ocean or respired back to CO2 and the release of other greenhouse gases; e.g., N2O. Central role played by pelagic fungi in the mycoloop and microbial loop.jpg
The central role played by pelagic fungi, both parasitic and saprotrophic in the mycoloop, and saprotrophic fungi as active contributors to the microbial loop. The activity of heterotrophic microbes, including pelagic fungi, has far-reaching global implications for fisheries (i.e., the amount of carbon that will ultimately flow to higher trophic levels) and climate change (i.e., the amount of carbon that will be sequestered in the ocean or respired back to CO2 and the release of other greenhouse gases; e.g., N2O.

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Parasites and food webs

Parasites are rarely included in food web studies, although they can strongly alter trophic interactions. In aquatic ecosystems, poorly grazed cyanobacteria often dominate phytoplankton communities, leading to the decoupling of primary and secondary production. [11] Advances in food web theory and modelling have contributed to understanding of the network of feeding relationships in ecological communities. Still, they often fail to explain processes observed in natural systems. [42] One reason for this is that most food web studies do not incorporate what is perhaps the most common trophic interaction - parasitism. [4] Despite their ubiquity, parasites are usually overlooked because of their cryptic nature, the difficulties in quantifying their effects, and their assumed low biomass. [5] However, they can account for greater biomass than predators [6] and participate in the majority of trophic links. [7] Parasites can modulate trophic flows in a number of ways. They can drive reductions in host biomass, not only by increasing host mortality rates, but also by influencing growth, fecundity, nutritional status, susceptibility to predation, or behaviour. [8] While their role as consumers is better known, parasites can also be prey for other organisms. They can be consumed together with their host (i.e. concomitant predation) or as free living life stages. Given the enormous reproductive output of parasites, free living infecting stages potentially constitute a significant nutrient source and can account for a substantial transfer of material and energy to higher trophic levels. [9] [11]

The efficiency of energy and material entry into the food web is largely determined by the trophic coupling between primary and secondary production. In aquatic pelagic ecosystems, primary production is often dominated by cyanobacteria. Promoted by eutrophication and global warming [43] [44] cyanobacteria often develop into blooms that severely disrupt ecosystem functioning and raise health concerns due to the production of diverse toxins [45] [46] Cyanobacteria display high resistance to grazing, which often leads to the decoupling of primary and secondary production and inefficient carbon transfer to zooplankton. [47] The inability of zooplankton to exert effective top-down control on cyanobacterial populations has traditionally been linked to the poor edibility of cyanobacteria with colonial or filamentous morphologies, the production of toxic metabolites, and their low nutritional value. [48] [11] [10]

Relation to the microbial carbon pump

Roles of fungi in the marine carbon cycle Roles of fungi in the marine carbon cycle by processing phytoplankton-derived organic matter. Parasitic fungi, as well as saprotrophic fungi, directly assimilate phytoplankton organic carbon. By releasing zoospores, the fungi bridge the trophic linkage to zooplankton, known as the mycoloop. By modifying the particulate and dissolved organic carbon, they can affect bacteria and the microbial loop. These processes may modify marine snow chemical composition and the subsequent functioning of the biological carbon pump. Roles of fungi in the marine carbon cycle.jpg
Roles of fungi in the marine carbon cycle
Roles of fungi in the marine carbon cycle by processing phytoplankton-derived organic matter. Parasitic fungi, as well as saprotrophic fungi, directly assimilate phytoplankton organic carbon. By releasing zoospores, the fungi bridge the trophic linkage to zooplankton, known as the mycoloop. By modifying the particulate and dissolved organic carbon, they can affect bacteria and the microbial loop. These processes may modify marine snow chemical composition and the subsequent functioning of the biological carbon pump.

Marine microorganisms make up around 70% of the total marine biomass [51] and are involved in complex functional and phylogenetic networks with all three organismal domains of life and viruses. [52] They harbor a set of genes responsible for driving major redox reactions that are crucial for controlling the remineralization of organic material. [53] Most of the research on the role of microbes in the oceanic nutrient cycling has focused on prokaryotes. Little is known on the role of pelagic fungi in the cycling of organic matter in the ocean despite fungi being recognized as key elements in remineralizing nutrients and degrading organic matter in the terrestrial and freshwater environment. [54] However, recent studies revealed that pelagic fungi were found to dominate the microbial biomass in deep-sea marine snow [55] and exhibited biomass concentrations similar to that of prokaryotes during phytoplankton blooms. [56] Moreover, by infecting inedible phytoplankton, parasitic fungi are suggested to act as trophic bridge via the fungal shunt by producing zoospores that are consumed by zooplankton, a process defined as the "mycoloop". [1] [57] [58] [59]

Recent evidence also indicates that pelagic fungi play a potentially important role in the marine carbon cycle. [60] [61] [62] A global-ocean scale multiomics study reported a widespread and active role of fungi in degrading carbohydrates by studying the diversity and expression of carbohydrate-active enzymes phylogenetically affiliated to fungi. [62] [59]

While the microbial carbon pump focuses on microbes transforming dissolved organic carbon (DOC) into refractory forms for long-term carbon sequestration in the deep ocean, the mycoloop is a food web process that transfers carbon and nutrients to higher trophic levels in surface waters. Both processes involve microbes (bacteria in microbial carbon pump, fungi in mycoloop) and contribute to carbon cycling, but the mycoloop emphasizes trophic interactions rather than long-term storage. [39] [63] [58]

References

  1. 1 2 Kagami, Maiko; von Elert, Eric; Ibelings, Bas W; de Bruin, Arnout; Van Donk, Ellen (22 June 2007). "The parasitic chytrid, Zygorhizidium , facilitates the growth of the cladoceran zooplankter, Daphnia , in cultures of the inedible alga, Asterionella". Proceedings of the Royal Society B: Biological Sciences. 274 (1617): 1561–1566. doi: 10.1098/rspb.2007.0425 . ISSN   0962-8452. PMC   2176168 . PMID   17439852.
  2. ↑ Grami, Boutheina; Rasconi, Serena; Niquil, Nathalie; Jobard, Marlène; Saint-Béat, Blanche; Sime-Ngando, Télesphore (22 August 2011). "Functional Effects of Parasites on Food Web Properties during the Spring Diatom Bloom in Lake Pavin: A Linear Inverse Modeling Analysis". PLOS ONE. 6 (8) e23273. Bibcode:2011PLoSO...623273G. doi: 10.1371/journal.pone.0023273 . ISSN   1932-6203. PMC   3161741 . PMID   21887240. CC-BY icon.svg Modified text was copied from this source, which is available under a Creative Commons Attribution 3.0 International License.
  3. 1 2 3 4 Danz, August; Quandt, C. Alisha (30 October 2023). "A review of the taxonomic diversity, host–parasite interactions, and experimental research on chytrids that parasitize diatoms". Frontiers in Microbiology. 14 1281648. doi: 10.3389/fmicb.2023.1281648 . ISSN   1664-302X. PMC   10643281 . PMID   38029223. CC-BY icon.svg Modified text was copied from this source, which is available under a Creative Commons Attribution 3.0 International License.
  4. 1 2 Marcogliese, David J.; Cone, David K. (1997). "Food webs: a plea for parasites" . Trends in Ecology & Evolution. 12 (8): 320–325. Bibcode:1997TEcoE..12..320M. doi:10.1016/S0169-5347(97)01080-X. PMID   21238094 . Retrieved 18 August 2025.
  5. 1 2 Lafferty, Kevin D.; Allesina, Stefano; Arim, Matias; Briggs, Cherie J.; De Leo, Giulio; Dobson, Andrew P.; Dunne, Jennifer A.; Johnson, Pieter T. J.; Kuris, Armand M.; Marcogliese, David J.; Martinez, Neo D.; Memmott, Jane; Marquet, Pablo A.; McLaughlin, John P.; Mordecai, Erin A.; Pascual, Mercedes; Poulin, Robert; Thieltges, David W. (2008). "Parasites in food webs: the ultimate missing links". Ecology Letters. 11 (6): 533–546. Bibcode:2008EcolL..11..533L. doi: 10.1111/j.1461-0248.2008.01174.x . ISSN   1461-023X. PMC   2408649 . PMID   18462196.
  6. 1 2 Kuris, Armand M.; Hechinger, Ryan F.; Shaw, Jenny C.; Whitney, Kathleen L.; Aguirre-Macedo, Leopoldina; Boch, Charlie A.; Dobson, Andrew P.; Dunham, Eleca J.; Fredensborg, Brian L.; Huspeni, Todd C.; Lorda, Julio; Mababa, Luzviminda; Mancini, Frank T.; Mora, Adrienne B.; Pickering, Maria; Talhouk, Nadia L.; Torchin, Mark E.; Lafferty, Kevin D. (24 July 2008). "Ecosystem energetic implications of parasite and free-living biomass in three estuaries" . Nature. 454 (7203): 515–518. Bibcode:2008Natur.454..515K. doi:10.1038/nature06970. ISSN   0028-0836. PMID   18650923 . Retrieved 18 August 2025.
  7. 1 2 Amundsen, Per-Arne; Lafferty, Kevin D.; Knudsen, Rune; Primicerio, Raul; Klemetsen, Anders; Kuris, Armand M. (2009). "Food web topology and parasites in the pelagic zone of a subarctic lake". Journal of Animal Ecology. 78 (3): 563–572. Bibcode:2009JAnEc..78..563A. doi: 10.1111/j.1365-2656.2008.01518.x . ISSN   0021-8790. PMID   19175443.
  8. 1 2 Selakovic, Sanja; de Ruiter, Peter C.; Heesterbeek, Hans (22 February 2014). "Infectious disease agents mediate interaction in food webs and ecosystems". Proceedings of the Royal Society B: Biological Sciences. 281 (1777) 20132709. doi: 10.1098/rspb.2013.2709 . ISSN   0962-8452. PMC   3896020 . PMID   24403336.
  9. 1 2 Johnson, Pieter T.J.; Dobson, Andrew; Lafferty, Kevin D.; Marcogliese, David J.; Memmott, Jane; Orlofske, Sarah A.; Poulin, Robert; Thieltges, David W. (2010). "When parasites become prey: ecological and epidemiological significance of eating parasites" . Trends in Ecology & Evolution. 25 (6): 362–371. Bibcode:2010TEcoE..25..362J. doi:10.1016/j.tree.2010.01.005. PMID   20185202 . Retrieved 18 August 2025.
  10. 1 2 Thieltges, David W.; Amundsen, Per-Arne; Hechinger, Ryan F.; Johnson, Pieter T. J.; Lafferty, Kevin D.; Mouritsen, Kim N.; Preston, Daniel L.; Reise, Karsten; Zander, C. Dieter; Poulin, Robert (2013). "Parasites as prey in aquatic food webs: implications for predator infection and parasite transmission" (PDF). Oikos. 122 (10): 1473–1482. Bibcode:2013Oikos.122.1473T. doi: 10.1111/j.1600-0706.2013.00243.x . hdl:10037/11842. ISSN   0030-1299 . Retrieved 18 August 2025.
  11. 1 2 3 4 Agha, Ramsy; Saebelfeld, Manja; Manthey, Christin; Rohrlack, Thomas; Wolinska, Justyna (13 October 2016). "Chytrid parasitism facilitates trophic transfer between bloom-forming cyanobacteria and zooplankton (Daphnia)" (PDF). Scientific Reports. 6 (1) 35039. Bibcode:2016NatSR...635039A. doi: 10.1038/srep35039 . ISSN   2045-2322. PMC   5062065 . PMID   27733762 . Retrieved 12 August 2025. CC-BY icon.svg Modified text was copied from this source, which is available under a Creative Commons Attribution 3.0 International License.
  12. ↑ Barr, D. J. S. (2001). "Chytridiomycota". Systematics and Evolution. Berlin, Heidelberg: Springer Berlin Heidelberg. pp. 93–112. doi:10.1007/978-3-662-10376-0_5. ISBN   978-3-642-08193-4 . Retrieved 27 August 2025.
  13. ↑ Gleason, Frank H.; Scholz, Bettina; Jephcott, Thomas G.; van Ogtrop, Floris F.; Henderson, Linda; Lilje, Osu; Kittelmann, Sandra; Macarthur, Deborah J. (10 March 2017). "Key Ecological Roles for Zoosporic True Fungi in Aquatic Habitats". Microbiology Spectrum. 5 (2) 5.2.17. doi:10.1128/microbiolspec.FUNK-0038-2016. ISSN   2165-0497. PMC   11687468 . PMID   28361735.
  14. ↑ Berger, Lee; Speare, Rick; Daszak, Peter; Green, D. Earl; Cunningham, Andrew A.; Goggin, C. Louise; Slocombe, Ron; Ragan, Mark A.; Hyatt, Alex D.; McDonald, Keith R.; Hines, Harry B.; Lips, Karen R.; Marantelli, Gerry; Parkes, Helen (21 July 1998). "Chytridiomycosis causes amphibian mortality associated with population declines in the rain forests of Australia and Central America". Proceedings of the National Academy of Sciences. 95 (15): 9031–9036. Bibcode:1998PNAS...95.9031B. doi: 10.1073/pnas.95.15.9031 . ISSN   0027-8424. PMC   21197 . PMID   9671799.
  15. ↑ Fisher, Matthew C.; Garner, Trenton W.J.; Walker, Susan F. (1 October 2009). "Global Emergence of Batrachochytrium dendrobatidis and Amphibian Chytridiomycosis in Space, Time, and Host" . Annual Review of Microbiology. 63 (1): 291–310. doi:10.1146/annurev.micro.091208.073435. ISSN   0066-4227. PMID   19575560 . Retrieved 27 August 2025.
  16. 1 2 Sparrow, F.K. (1960). Aquatic Phycomyete (2nd ed.). Ann Arbor, MI: The University of Michigan Press.
  17. ↑ Hibbett; et al. (2007). "A higher-level phylogenetic classification of the Fungi". Mycologia. 111 (5): 509–547. doi:10.1016/j.mycres.2007.03.004. PMID   17572334. S2CID   4686378.
  18. 1 2 Frenken, T., Alacid, E., Berger, S.A., Bourne, E.C., Gerphagnon, M., Grossart, H.P., Gsell, A.S., Ibelings, B.W., Kagami, M., Küpper, F.C. and Letcher, P.M. (2017) "Integrating chytrid fungal parasites into plankton ecology: research gaps and needs". Environmental microbiology, 19(10): 3802–3822. doi : 10.1111/1462-2920.13827. CC-BY icon.svg Modified text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  19. ↑ Kilias, Estelle S.; Junges, Leandro; Šupraha, Luka; Leonard, Guy; Metfies, Katja; Richards, Thomas A. (2020). "Chytrid fungi distribution and co-occurrence with diatoms correlate with sea ice melt in the Arctic Ocean". Communications Biology. 3 (1): 183. doi:10.1038/s42003-020-0891-7. PMC   7174370 . PMID   32317738. S2CID   216033140. CC-BY icon.svg Modified text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  20. ↑ Maier, Michelle A.; Peterson, Tawnya D. (2014). "Observations of a Diatom Chytrid Parasite in the Lower Columbia River" . Northwest Science. 88 (3): 234–245. Bibcode:2014NWSci..88..234M. doi:10.3955/046.088.0306. ISSN   0029-344X . Retrieved 8 August 2025.
  21. ↑ Jones EG, Hyde KD, Pang KL, eds. (27 August 2014). Freshwater Fungi: and Fungal-like Organisms. Walter de Gruyter GmbH & Co KG. ISBN   978-3-11-033348-0.
  22. ↑ Jones EG, Hyde KD, Pang KL (31 August 2012). Marine Fungi: and Fungal-like Organisms. Walter de Gruyter. ISBN   978-3-11-026406-7.
  23. 1 2 Sen, Kalymani; Sen, Biswarup; Wang, Guangyi (8 May 2022). "Diversity, Abundance, and Ecological Roles of Planktonic Fungi in Marine Environments". Journal of Fungi. 8 (5): 491. doi: 10.3390/jof8050491 . PMC   9147564 . PMID   35628747.
  24. ↑ Wang X, Singh P, Gao Z, Zhang X, Johnson ZI, Wang G (3 July 2014). "Distribution and diversity of planktonic fungi in the West Pacific Warm Pool". PLOS ONE. 9 (7) e101523. Bibcode:2014PLoSO...9j1523W. doi: 10.1371/journal.pone.0101523 . PMC   4081592 . PMID   24992154.
  25. ↑ Raghukumar C, ed. (2012). Biology of Marine Fungi. Progress in Molecular and Subcellular Biology. Vol. 53. doi:10.1007/978-3-642-23342-5. ISBN   978-3-642-23341-8. ISSN   0079-6484. S2CID   39378040.
  26. ↑ Damare S, Raghukumar C (July 2008). "Fungi and macroaggregation in deep-sea sediments". Microbial Ecology. 56 (1): 168–177. Bibcode:2008MicEc..56..168D. doi:10.1007/s00248-007-9334-y. PMID   17994287. S2CID   21288251.
  27. ↑ Cudowski, A.; Pietryczuk, A. (2020). "Biodiversity of mycoplankton in the profile of eutrophic lakes with varying water quality" . Fungal Ecology. 48 100978. Bibcode:2020FunE...4800978C. doi:10.1016/j.funeco.2020.100978 . Retrieved 26 August 2025.
  28. ↑ Sen, Kalyani; Sen, Biswarup; Wang, Guangyi (8 May 2022). "Diversity, Abundance, and Ecological Roles of Planktonic Fungi in Marine Environments". Journal of Fungi. 8 (5): 491. doi: 10.3390/jof8050491 . ISSN   2309-608X. PMC   9147564 . PMID   35628747.
  29. ↑ Kubanek J, Jensen PR, Keifer PA, Sullards MC, Collins DO, Fenical W (June 2003). "Seaweed resistance to microbial attack: a targeted chemical defense against marine fungi". Proceedings of the National Academy of Sciences of the United States of America. 100 (12): 6916–6921. Bibcode:2003PNAS..100.6916K. doi: 10.1073/pnas.1131855100 . PMC   165804 . PMID   12756301.
  30. ↑ Jobard M, Rasconi S, Sime-Ngando T (1 June 2010). "Diversity and functions of microscopic fungi: a missing component in pelagic food webs". Aquatic Sciences. 72 (3): 255–268. Bibcode:2010AqSci..72..255J. doi:10.1007/s00027-010-0133-z. ISSN   1420-9055. S2CID   36789070.
  31. ↑ Gao Z, Johnson ZI, Wang G (January 2010). "Molecular characterization of the spatial diversity and novel lineages of mycoplankton in Hawaiian coastal waters". The ISME Journal. 4 (1): 111–120. Bibcode:2010ISMEJ...4..111G. doi: 10.1038/ismej.2009.87 . PMID   19641535. S2CID   2395339.
  32. ↑ Panzer K, Yilmaz P, Weiß M, Reich L, Richter M, Wiese J, et al. (30 July 2015). "Identification of Habitat-Specific Biomes of Aquatic Fungal Communities Using a Comprehensive Nearly Full-Length 18S rRNA Dataset Enriched with Contextual Data". PLOS ONE. 10 (7) e0134377. Bibcode:2015PLoSO..1034377P. doi: 10.1371/journal.pone.0134377 . PMC   4520555 . PMID   26226014.
  33. ↑ "First record of flamentous fungi in the coastal upwelling ecosystem off central Chile". Gayana (Concepción). 68 (2). 2004. doi: 10.4067/s0717-65382004000200001 . ISSN   0717-6538.
  34. ↑ Sridhar KR (January 2009). "Aquatic fungi – Are they planktonic?". Plankton Dynamics of Indian Waters. Jaipur, India: Pratiksha Publications. pp. 133–148.
  35. 1 2 Plankton Manifesto Ocean Stewardship Coalition of the United Nations Global Compact, published September 2024.
  36. ↑ Kagami, M., Miki, T. and Takimoto, G. (2014) "Mycoloop: chytrids in aquatic food webs". Frontiers in microbiology, 5: 166. doi : 10.3389/fmicb.2014.00166 . CC-BY icon.svg Modified text was copied from this source, which is available under a Creative Commons Attribution 3.0 International License.
  37. ↑ Amend, A., Burgaud, G., Cunliffe, M., Edgcomb, V.P., Ettinger, C.L., Gutiérrez, M.H., Heitman, J., Hom, E.F., Ianiri, G., Jones, A.C. and Kagami, M. (2019) "Fungi in the marine environment: Open questions and unsolved problems". MBio, 10(2): e01189-18. doi : 10.1128/mBio.01189-18. CC-BY icon.svg Modified text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  38. ↑ Gutiérrez, Marcelo H.; Jara, Ana M.; Pantoja, Silvio (May 2016). "Fungal parasites infect marine diatoms in the upwelling ecosystem of the Humboldt current system off central Chile" . Environmental Microbiology. 18 (5): 1646–1653. Bibcode:2016EnvMi..18.1646G. doi:10.1111/1462-2920.13257. hdl: 10533/148260 . ISSN   1462-2912. PMID   26914416.
  39. 1 2 3 Kagami, M., Miki, T. and Takimoto, G. (2014) "Mycoloop: chytrids in aquatic food webs". Frontiers in microbiology, 5: 166. doi : 10.3389/fmicb.2014.00166 . CC-BY icon.svg Modified text was copied from this source, which is available under a Creative Commons Attribution 3.0 International License.
  40. ↑ Barranco, V.S., Van der Meer, M.T., Kagami, M., Van den Wyngaert, S., Van de Waal, D.B., Van Donk, E. and Gsell, A.S. (2020( "Trophic position, elemental ratios and nitrogen transfer in a planktonic host–parasite–consumer food chain including a fungal parasite". Oecologia: 1–14. doi : 10.1007/s00442-020-04721-w. CC-BY icon.svg Modified text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  41. ↑ Breyer, Eva; Baltar, Federico (2023). "The largely neglected ecological role of oceanic pelagic fungi". Trends in Ecology & Evolution. 38 (9): 870–888. Bibcode:2023TEcoE..38..870B. doi: 10.1016/j.tree.2023.05.002 . PMID   37246083 . Retrieved 27 July 2025.
  42. ↑ Brett, M T; Goldman, C R (23 July 1996). "A meta-analysis of the freshwater trophic cascade". Proceedings of the National Academy of Sciences. 93 (15): 7723–7726. Bibcode:1996PNAS...93.7723B. doi: 10.1073/pnas.93.15.7723 . ISSN   0027-8424. PMC   38814 . PMID   11607694.
  43. ↑ Markensten, Hampus; Moore, Karen; Persson, Irina (2010). "Simulated lake phytoplankton composition shifts toward cyanobacteria dominance in a future warmer climate" . Ecological Applications. 20 (3): 752–767. Bibcode:2010EcoAp..20..752M. doi:10.1890/08-2109.1. ISSN   1051-0761. PMID   20437961 . Retrieved 19 August 2025.
  44. ↑ Paerl, Hans W.; Huisman, Jef (2009). "Climate change: a catalyst for global expansion of harmful cyanobacterial blooms" . Environmental Microbiology Reports. 1 (1): 27–37. Bibcode:2009EnvMR...1...27P. doi:10.1111/j.1758-2229.2008.00004.x. ISSN   1758-2229. PMID   23765717 . Retrieved 19 August 2025.
  45. ↑ Codd, Geoffrey A.; Morrison, Louise F.; Metcalf, James S. (2005). "Cyanobacterial toxins: risk management for health protection" . Toxicology and Applied Pharmacology. 203 (3): 264–272. Bibcode:2005ToxAP.203..264C. doi:10.1016/j.taap.2004.02.016. PMID   15737680 . Retrieved 19 August 2025.
  46. ↑ Zillén, L.; Conley, D. J. (31 August 2010). "Hypoxia and cyanobacteria blooms - are they really natural features of the late Holocene history of the Baltic Sea?". Biogeosciences. 7 (8): 2567–2580. Bibcode:2010BGeo....7.2567Z. doi: 10.5194/bg-7-2567-2010 . ISSN   1726-4189.
  47. ↑ De Bernardi, R. d. & Giussani, G. Biomanipulation Tool for Water Management: Proceedings of an International Conference held in Amsterdam, 8–11 Aug 1989, pp. 29–41.
  48. ↑ Ger, Kemal Ali; Urrutia-Cordero, Pablo; Frost, Paul C.; Hansson, Lars-Anders; Sarnelle, Orlando; Wilson, Alan E.; Lürling, Miquel (2016). "The interaction between cyanobacteria and zooplankton in a more eutrophic world" . Harmful Algae. 54: 128–144. Bibcode:2016HAlga..54..128G. doi:10.1016/j.hal.2015.12.005. PMID   28073472 . Retrieved 19 August 2025.
  49. ↑ Amend, A., Burgaud, G., Cunliffe, M., Edgcomb, V.P., Ettinger, C.L., Gutiérrez, M.H., Heitman, J., Hom, E.F., Ianiri, G., Jones, A.C. and Kagami, M. (2019) "Fungi in the marine environment: Open questions and unsolved problems". MBio, 10(2): e01189-18. doi : 10.1128/mBio.01189-18. CC-BY icon.svg Modified text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  50. ↑ Gutierrez MH, Jara AM, Pantoja S (2016) "Fungal parasites infect marine diatoms in the upwelling ecosystem of the Humboldt current system off central Chile". Environ Microbiol, 18(5): 1646–1653. doi : 10.1111/1462-2920.13257.
  51. ↑ Bar-On, Yinon M.; Phillips, Rob; Milo, Ron (19 June 2018). "The biomass distribution on Earth" (PDF). Proceedings of the National Academy of Sciences. 115 (25): 6506–6511. Bibcode:2018PNAS..115.6506B. doi: 10.1073/pnas.1711842115 . ISSN   0027-8424. PMC   6016768 . PMID   29784790 . Retrieved 14 August 2025.
  52. ↑ Lima-Mendez, Gipsi; Faust, Karoline; Henry, Nicolas; Decelle, Johan; et al. (22 May 2015). "Determinants of community structure in the global plankton interactome" (PDF). Science. 348 (6237) 1262073. doi: 10.1126/science.1262073 . hdl:2078.1/231553. ISSN   0036-8075 . Retrieved 15 August 2025.
  53. ↑ Falkowski, Paul G.; Fenchel, Tom; Delong, Edward F. (23 May 2008). "The Microbial Engines That Drive Earth's Biogeochemical Cycles" (PDF). Science. 320 (5879): 1034–1039. Bibcode:2008Sci...320.1034F. doi: 10.1126/science.1153213 . ISSN   0036-8075. PMID   18497287 . Retrieved 15 August 2025.
  54. ↑ Grossart, Hans-Peter; Van den Wyngaert, Silke; Kagami, Maiko; Wurzbacher, Christian; et al. (2019). "Fungi in aquatic ecosystems". Nature Reviews Microbiology. 17 (6): 339–354. doi: 10.1038/s41579-019-0175-8 . hdl: 10669/81667 . ISSN   1740-1526. PMID   30872817 . Retrieved 15 August 2025.
  55. ↑ Bochdansky, Alexander B; Clouse, Melissa A; Herndl, Gerhard J (20 September 2016). "Eukaryotic microbes, principally fungi and labyrinthulomycetes, dominate biomass on bathypelagic marine snow" (PDF). The ISME Journal. 11 (2). Oxford University Press (OUP): 362–373. doi: 10.1038/ismej.2016.113 . ISSN   1751-7362. PMC   5270556 . PMID   27648811 . Retrieved 15 August 2025.
  56. ↑ Gutiérrez, M. H.; Pantoja, S.; Tejos, E.; Quiñones, R. A. (2011). "The role of fungi in processing marine organic matter in the upwelling ecosystem off Chile" . Marine Biology. 158 (1): 205–219. Bibcode:2011MarBi.158..205G. doi:10.1007/s00227-010-1552-z. ISSN   0025-3162 . Retrieved 15 August 2025.
  57. ↑ Cleary, Alison C.; Søreide, Janne E.; Freese, Daniela; Niehoff, Barbara; Gabrielsen, Tove M. (1 October 2017). "Feeding by Calanus glacialis in a high arctic fjord: potential seasonal importance of alternative prey" (PDF). ICES Journal of Marine Science. 74 (7): 1937–1946. doi: 10.1093/icesjms/fsx106 . ISSN   1054-3139 . Retrieved 15 August 2025.
  58. 1 2 Klawonn, Isabell; Van den Wyngaert, Silke; Parada, Alma E.; Arandia-Gorostidi, Nestor; et al. (8 June 2021). "Characterizing the "fungal shunt": Parasitic fungi on diatoms affect carbon flow and bacterial communities in aquatic microbial food webs". Proceedings of the National Academy of Sciences. 118 (23) e2102225118. Bibcode:2021PNAS..11802225K. doi: 10.1073/pnas.2102225118 . ISSN   0027-8424. PMC   8201943 . PMID   34074785.
  59. 1 2 Breyer, Eva; Zhao, Zihao; Herndl, Gerhard J.; Baltar, Federico (1 September 2022). "Global contribution of pelagic fungi to protein degradation in the ocean". Microbiome. 10 (1) 143. doi: 10.1186/s40168-022-01329-5 . ISSN   2049-2618. PMC   9434897 . PMID   36050758. CC-BY icon.svg Modified text was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
  60. ↑ Morales, Sergio E.; Biswas, Ambarish; Herndl, Gerhard J.; Baltar, Federico (19 March 2019). "Global Structuring of Phylogenetic and Functional Diversity of Pelagic Fungi by Depth and Temperature". Frontiers in Marine Science. 6 131. Bibcode:2019FrMaS...6..131M. doi: 10.3389/fmars.2019.00131 . ISSN   2296-7745.
  61. ↑ Chrismas, Nathan; Cunliffe, Michael (1 September 2020). "Depth-dependent mycoplankton glycoside hydrolase gene activity in the open ocean—evidence from the Tara Oceans eukaryote metatranscriptomes" (PDF). The ISME Journal. 14 (9): 2361–2365. Bibcode:2020ISMEJ..14.2361C. doi: 10.1038/s41396-020-0687-2 . ISSN   1751-7362. PMC   7608184 . PMID   32494052 . Retrieved 15 August 2025.
  62. 1 2 Baltar, Federico; Zhao, Zihao; Herndl, Gerhard J. (2021). "Potential and expression of carbohydrate utilization by marine fungi in the global ocean". Microbiome. 9 (1) 106. doi: 10.1186/s40168-021-01063-4 . ISSN   2049-2618. PMC   8114511 . PMID   33975640.
  63. ↑ Chen, Ming; Gao, Honghui; Zhang, Jimin (2024). "Mycoloop: Modeling phytoplankton–chytrid–zooplankton interactions in aquatic food webs" . Mathematical Biosciences. 368 109134. doi:10.1016/j.mbs.2023.109134. PMID   38158013 . Retrieved 10 August 2025.
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