Paramecium bursaria

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Paramecium bursaria
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Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Clade: Diaphoretickes
Clade: SAR
Clade: Alveolata
Phylum: Ciliophora
Class: Oligohymenophorea
Order: Peniculida
Family: Parameciidae
Genus: Paramecium
Species:
P. bursaria
Binomial name
Paramecium bursaria
(Ehr., 1831)

Background

Paramecium bursaria is a species of ciliate found in marine and brackish waters. [1] It has a mutualistic endosymbiotic relationship with green algae called Zoochlorella . About 700 Chlorella cells live inside the protist's cytoplasm and provide it with food, while the Paramecium provides the algae with movement and protection. [2] P. bursaria is 80-150 μm long, with a wide oral groove, two contractile vacuoles, and a single micronucleus as well as a single macronucleus. P. bursaria is the only species of Paramecium that forms symbiotic relationships with algae, and it is often used in biology classrooms both as an example of a protozoan and also as an example of symbiosis.

Contents

A transcriptome sequence is determined. [3]

Core principle of endosymbiosis

Paramecium bursaria harbors approximately 700 cells of zoochlorellae (green algae) from the genera Chlorella or Micractinium under its cell cortex, forming endosymbionts. [4] [5] The core principle of these endosymbionts is nutrition, where the host obtains nutrients through phagotrophy by engulfing cells or particles, including Chlorella, which are digested in the digestive vacuole (DV). However, some Chlorella cells can resist this digestive process and become enclosed in a distinct vacuole, known as the perialgal vacuole (PV), formed by the host. This vacuole prevents lysosomal fusion, allowing the algae to survive and establish symbiosis. [6] The PV can be distinguished from the digestive vacuole (DV) by the number and distribution of particles, exhibiting minimal endocytosis or exocytosis activity.

Moreover, the mechanism of the symbiotic relationship between Paramecium bursaria and Chlorella, based on comparative genome analysis, suggests that the host provides glutamine and magnesium. Chlorella utilizes glutamine as a nitrogen source and magnesium to support chlorophyll-based photosynthesis. In return, Chlorella within the perialgal vacuole (PV) provides the host with photosynthetic products such as fructose, maltose, and oxygen. [7]

It is also worth highlighting how the host regulates nutrient exchange in this photosynthetic symbiosis. Andrew et al. (2016) developed a mathematical model, using the framework of general symbiotic relationships, to describe the mechanism of this interaction. During vertical transmission of symbionts through cytokinesis, daughter cells receive an equal number of symbionts as their parent cells. However, during horizontal transmission, the daughter cells acquire only half the symbionts. The host's growth rate is influenced by its nutritional state, which depends on the efficiency of nutrient exchange with the symbionts. By managing this exchange, the host can optimize its growth and the overall benefits of the symbiosis.

Focusing on light levels as an environmental condition, the study investigated how the host regulates this mechanism. The results show that as light levels increase, there is a reduction in the symbiont population. This regulation is crucial for maintaining a stable symbiont population and preventing parasitism, which could occur if the symbionts were to overgrow and harm the host. [6]

Phylogenetic analysis

In terms of phylogenetic analysis, a study by Christian et al. (2022) collected 48 strains of Paramecium bursaria from around the world and analyzed their phylogenetic lineages using SSU and ITS rDNA sequences. The results revealed five highly supported lineages corresponding to the syngens (R1–R5), which are genetically distinct but reproductively isolated among the P. bursaria strains sharing synapomorphic traits. This study also confirmed the presence of syngens (R1–R5) based on the secondary structures of the SSU and ITS regions, as well as synapomorphies in the amino acid sequences of the COI gene, aiding in syngen identification. These findings enabled the clear assignment of existing P. bursaria entries in GenBank to specific syngens.

In addition, the study examined the morphology and phenotypic plasticity by selecting at least one strain from each lineage: P. protobursaria from syngen R1, P. deuterobursaria from syngen R2, P. tritobursaria from syngen R3, P. tetratobursaria from syngen R4, and P. pentobursaria from syngen R5. In terms of cell length and width, differences among the syngens were not highly significant, although syngen R5 was the longest, followed by R4, R3, R2, and R1, respectively. All syngens were ellipsoidal to broadly ellipsoidal and dorsoventrally flattened in cell shape.

Regarding their symbiotic associations, syngens R1 and R2 formed symbioses with Micractinium conductrix, whereas R3 and R5 associated with Chlorella variabilis. Syngen R4 displayed the ability to host all three algal strains: Micractinium conductrix, Chlorella variabilis, and Chlorella vulgaris. [8]

Genomic analysis

Moreover, a combination of long-read sequencing (PacBio) and short-read sequencing (Illumina) can be used to assemble a high-quality, near-complete macronuclear genome of P. bursaria, providing insights into the mechanism of endosymbiosis. The genome size of P. bursaria is much smaller than that of other ciliate species but is comparable to P. caudatum.

Additionally, the standard core genes in the assembled genome of P. bursaria (94.8%) are similar to those of other well-studied ciliate genomes, such as T. thermophila (89.9%), O. trifallax (93.1%), and P. tetraurelia (92.7%). [7]

Related Research Articles

<span class="mw-page-title-main">Endosymbiont</span> Organism that lives within the body or cells of another organism

An endosymbiont or endobiont is an organism that lives within the body or cells of another organism. Typically the two organisms are in a mutualistic relationship. Examples are nitrogen-fixing bacteria, which live in the root nodules of legumes, single-cell algae inside reef-building corals, and bacterial endosymbionts that provide essential nutrients to insects.

<span class="mw-page-title-main">Symbiosis</span> Close, long-term biological interaction between distinct organisms (usually species)

Symbiosis is any type of a close and long-term biological interaction, between two organisms of different species. The two organisms, termed symbionts, can be either in a mutualistic, a commensalistic, or a parasitic relationship. In 1879, Heinrich Anton de Bary defined symbiosis as "the living together of unlike organisms".

<i>Paramecium</i> Genus of unicellular ciliates, commonly studied as a representative of the ciliate group

Paramecium is a genus of eukaryotic, unicellular ciliates, widespread in freshwater, brackish, and marine environments. Paramecia are often abundant in stagnant basins and ponds. Because some species are readily cultivated and easily induced to conjugate and divide, they have been widely used in classrooms and laboratories to study biological processes. Paramecium species are commonly studied as model organisms of the ciliate group and have been characterized as the "white rats" of the phylum Ciliophora.

<span class="mw-page-title-main">Zooxanthellae</span> Dinoflagellates in symbiosis with coral, jellyfish and nudibranchs

Zooxanthellae is a colloquial term for single-celled dinoflagellates that are able to live in symbiosis with diverse marine invertebrates including demosponges, corals, jellyfish, and nudibranchs. Most known zooxanthellae are in the genus Symbiodinium, but some are known from the genus Amphidinium, and other taxa, as yet unidentified, may have similar endosymbiont affinities. "Zooxanthella" was originally a genus name given in 1881 by Karl Brandt to Zooxanthella nutricula which has been placed in the Peridiniales. Another group of unicellular eukaryotes that partake in similar endosymbiotic relationships in both marine and freshwater habitats are green algae zoochlorellae.

Zoochlorella is a coloquial term for any green algae that lives symbiotically within the body of an aquatic invertebrate animal or a protozoan.

<i>Stentor</i> (ciliate) Genus of single-celled organisms

Stentor, sometimes called trumpet animalcules, are a genus of filter-feeding, heterotrophic ciliates, representative of the heterotrichs. They are usually horn-shaped, and reach lengths of two millimeters; as such, they are among the largest known extant unicellular organisms. They reproduce asexually through binary fission.

<span class="mw-page-title-main">Bacteriocyte</span> Specialized cell containing endosymbionts

A bacteriocyte, also known as a mycetocyte, is a specialized adipocyte found primarily in certain insects such as aphids, tsetse flies, German cockroaches, weevils, and ants. These cells contain endosymbiotic organisms such as bacteria and fungi, which provide essential amino acids and other chemicals to their host. Bacteriocytes may aggregate into a specialized organ called the bacteriome.

<i>Mixotricha paradoxa</i> Species of protozoan

Mixotricha paradoxa is a species of protozoan that lives inside the gut of the Australian termite species Mastotermes darwiniensis.

Symbiotic bacteria are bacteria living in symbiosis with another organism or each other. For example, rhizobia living in root nodules of legumes provide nitrogen fixing activity for these plants.

Phycodnaviridae is a family of large (100–560 kb) double-stranded DNA viruses that infect marine or freshwater eukaryotic algae. Viruses within this family have a similar morphology, with an icosahedral capsid. As of 2014, there were 33 species in this family, divided among 6 genera. This family belongs to a super-group of large viruses known as nucleocytoplasmic large DNA viruses. Evidence was published in 2014 suggesting that specific strains of Phycodnaviridae might infect humans rather than just algal species, as was previously believed. Most genera under this family enter the host cell by cell receptor endocytosis and replicate in the nucleus. Phycodnaviridae play important ecological roles by regulating the growth and productivity of their algal hosts. Algal species such Heterosigma akashiwo and the genus Chrysochromulina can form dense blooms which can be damaging to fisheries, resulting in losses in the aquaculture industry. Heterosigma akashiwo virus (HaV) has been suggested for use as a microbial agent to prevent the recurrence of toxic red tides produced by this algal species. Phycodnaviridae cause death and lysis of freshwater and marine algal species, liberating organic carbon, nitrogen and phosphorus into the water, providing nutrients for the microbial loop.

A mixotroph is an organism that uses a mix of different sources of energy and carbon, instead of having a single trophic mode, on the continuum from complete autotrophy to complete heterotrophy. It is estimated that mixotrophs comprise more than half of all microscopic plankton. There are two types of eukaryotic mixotrophs. There are those with their own chloroplasts - including those with endosymbionts providing the chloroplasts. And there are those that acquire them through kleptoplasty, or through symbiotic associations with prey, or through 'enslavement' of the prey's organelles.

<i>Climacostomum</i> Genus of single-celled organisms

Climacostomum is a genus of unicellular ciliates, belonging to the class Heterotrichea.

Chlorovirus, also known as Chlorella virus, is a genus of giant double-stranded DNA viruses, in the family Phycodnaviridae. This genus is found globally in freshwater environments where freshwater microscopic algae serve as natural hosts. There are 19 species in this genus.

<span class="mw-page-title-main">Marine microbial symbiosis</span>

Microbial symbiosis in marine animals was not discovered until 1981. In the time following, symbiotic relationships between marine invertebrates and chemoautotrophic bacteria have been found in a variety of ecosystems, ranging from shallow coastal waters to deep-sea hydrothermal vents. Symbiosis is a way for marine organisms to find creative ways to survive in a very dynamic environment. They are different in relation to how dependent the organisms are on each other or how they are associated. It is also considered a selective force behind evolution in some scientific aspects. The symbiotic relationships of organisms has the ability to change behavior, morphology and metabolic pathways. With increased recognition and research, new terminology also arises, such as holobiont, which the relationship between a host and its symbionts as one grouping. Many scientists will look at the hologenome, which is the combined genetic information of the host and its symbionts. These terms are more commonly used to describe microbial symbionts.

<span class="mw-page-title-main">Armophorea</span> Class of single-celled organisms

Armophorea is a class of ciliates in the subphylum Intramacronucleata. . It was first resolved in 2004 and comprises three orders: Metopida, Clevelandellida, and Armophorida. Previously members of this class were thought to be heterotrichs because of similarities in morphology, most notably a characteristic dense arrangement of cilia surrounding their oral structures. However, the development of genetic tools and subsequent incorporation of DNA sequence information has led to major revisions in the evolutionary relationships of many protists, including ciliates. Metopids, clevelandellids, and armophorids were grouped into this class based on similarities in their small subunit rRNA sequences, making them one of two so-called "riboclasses" of ciliates, however, recent analyses suggest that Armophorida may not be related to the other two orders.

<i>Hydra viridissima</i> Species of hydrozoan

Hydra viridissima is a species of cnidarian which is commonly found in still or slow-moving freshwater in the Northern temperate zone. Hydra viridissima is commonly called green hydra due to its coloration, which is due to the symbiotic green algae Chlorella vulgaris which live within its body. These creatures are typically 10 mm long and have tentacles that are about half of their length. They are strictly carnivorous and typically feed on small crustaceans, insects and annelids. Hydra are normally sessile and live on aquatic vegetation. They secrete mucus to attach to substrate using their basal disc.

<span class="mw-page-title-main">Symbiosome</span>

A symbiosome is a specialised compartment in a host cell that houses an endosymbiont in a symbiotic relationship.

<span class="mw-page-title-main">Marine protists</span> Protists that live in saltwater or brackish water

Marine protists are defined by their habitat as protists that live in marine environments, that is, in the saltwater of seas or oceans or the brackish water of coastal estuaries. Life originated as marine single-celled prokaryotes and later evolved into more complex eukaryotes. Eukaryotes are the more developed life forms known as plants, animals, fungi and protists. Protists are the eukaryotes that cannot be classified as plants, fungi or animals. They are mostly single-celled and microscopic. The term protist came into use historically as a term of convenience for eukaryotes that cannot be strictly classified as plants, animals or fungi. They are not a part of modern cladistics because they are paraphyletic.

Novymonas esmeraldas is a protist and member of flagellated trypanosomatids. It is an obligate parasite in the gastrointestinal tract of a bug, and is in turn a host to symbiotic bacteria. It maintains strict mutualistic relationship with the bacteria as a sort of cell organelle (endosymbiont) so that it cannot lead an independent life without the bacteria. Its discovery in 2016 suggests that it is a good model in the evolution of prokaryotes into eukaryotes by symbiogenesis. The endosymbiotic bacterium was identified as member of the genus Pandoraea.

<i>Pseudoblepharisma</i> Genus of protozoans

Pseudoblepharisma is a genus of heterotrich ciliates inhabiting oxygen depleted freshwater habitats. Most sources report that it contains one species, Pseudoblepharisma tenue, but at least four have been seen in literature.

References

  1. Warren, A. (2019) Paramecium bursaria (Ehrenberg) Focker, 1836 WoRMS. Accessed: 16 October 2019.
  2. Kodama, Yuuki; Fujishima, Masahiro (2022). "Endosymbiotic Chlorella variabilis reduces mitochondrial number in the ciliate Paramecium bursaria". Scientific Reports. 12 (1): 8216. Bibcode:2022NatSR..12.8216K. doi:10.1038/s41598-022-12496-8. PMC   9151773 . PMID   35637201.
  3. Dohra H; et al. (2015). "Analysis of amino acid and codon usage in Paramecium bursaria". FEBS Lett. 589 (20 Pt B): 3113–3118. Bibcode:2015FEBSL.589.3113D. doi:10.1016/j.febslet.2015.08.033. hdl: 10297/9791 . PMID   26341535. S2CID   357977.
  4. Spanner, Christian; Darienko, Tatyana; Filker, Sabine; Sonntag, Bettina; Pröschold, Thomas (2022-10-27). "Morphological diversity and molecular phylogeny of five Paramecium bursaria (Alveolata, Ciliophora, Oligohymenophorea) syngens and the identification of their green algal endosymbionts". Scientific Reports. 12 (1): 18089. doi:10.1038/s41598-022-22284-z. ISSN   2045-2322. PMC   9613978 . PMID   36302793.
  5. Kodama, Yuuki; Fujishima, Masahiro (2012-07-01). "Characteristics of the Digestive Vacuole Membrane of the Alga-Bearing Ciliate Paramecium bursaria". Protist. 163 (4): 658–670. doi:10.1016/j.protis.2011.10.004. ISSN   1434-4610.
  6. 1 2 Dean, Andrew D.; Minter, Ewan J. A.; Sørensen, Megan E. S.; Lowe, Christopher D.; Cameron, Duncan D.; Brockhurst, Michael A.; Jamie Wood, A. (2016-09-21). "Host control and nutrient trading in a photosynthetic symbiosis". Journal of Theoretical Biology. Advances in Modelling Biological Evolution: Linking Mathematical Theories with Empirical Realities. 405: 82–93. doi:10.1016/j.jtbi.2016.02.021. ISSN   0022-5193.
  7. 1 2 He, Ming; Wang, Jinfeng; Fan, Xinpeng; Liu, Xiaohui; Shi, Wenyu; Huang, Ning; Zhao, Fangqing; Miao, Miao (2019-05-01). "Genetic basis for the establishment of endosymbiosis in Paramecium". The ISME Journal. 13 (5): 1360–1369. doi:10.1038/s41396-018-0341-4. ISSN   1751-7362. PMC   6474222 . PMID   30647459.
  8. Spanner, Christian; Darienko, Tatyana; Filker, Sabine; Sonntag, Bettina; Pröschold, Thomas (2022-10-27). "Morphological diversity and molecular phylogeny of five Paramecium bursaria (Alveolata, Ciliophora, Oligohymenophorea) syngens and the identification of their green algal endosymbionts". Scientific Reports. 12 (1): 18089. doi:10.1038/s41598-022-22284-z. ISSN   2045-2322. PMC   9613978 . PMID   36302793.

Further reading