Why does stentor belong to the phylum ciliophora




















The genus Neisseria and other representatives of the family Neisseriaceae were found in rather high abundancy Table 2. Neisseria species normally colonize the mucosal surface of mammals and rarely invade their host cells 56 , 75 , so their presence in association with stentors might reveal novel aspects of this bacterial genus biology. Importantly, all these bacteria were absent or, like Streptococcus sp. Numerous OTUs associated with bacteria occasionally appearing as dominant microbiome components were defined in several Stentor samples.

These bacteria belong either to presumably free-living genera like bacterium MNG7 or Xanthobacter sp. We suggest to consider such bacteria as transitory components of Stentor microbiome, which by chance manage to colonize some ciliate cells.

We do not have an idea of any microbiome function useful for the host ciliate. Still, there is a common feature for almost all major bacteria in Stentor microbiome. These bacteria are very likely prone to interact with eukaryots, as they belong to genera encompassing numerous commensals or even pathogens of animals.

When they appear freely in water, they may use their general skills to interact with available eukaryotic cells, namely, protists. There are two possible ways of bacterial persistence in association with protists: to stay attached on the surface of the host cell, or to resist digestion or even to escape from the food vacuoles to the host cytoplasm. Ciliates are rather big protists S. Bacteria propagating on ciliate cell surface have been described 77 , Sessile ciliates may colonize big areas of suitable substrates, and in this way bacterial community associated with ciliate cells may get stabilized in certain general environmental conditions.

At the same time, ciliates can swim quickly and use different taxis allowing them to select favorable conditions 79 , which would be beneficial also for the associated bacteria. Ciliate surface may also offer a shelter to bacteria, protecting them from phagocytosis by other microorganisms. It is known that bacteria have developed numerous strategies to avoid prey-selective grazing by protozoa, namely morphological changes, high speed motility, and production of antagonistic or toxic substances 80 , 81 , 82 , One of the most important defense mechanisms is formation of biofilms or aggregations, in which separate bacteria are less vulnerable for predators From this point of view, the ciliate cell surface allows bacteria to adhere, aggregate, form biofilms and co-regulate their living activities through a quorum sensing mechanism, which regulates gene expression as a response to fluctuations in bacterial population density Thus, even a small substrate may be substantial for propagation of certain bacteria in the environment.

To our best knowledge, no one have tried yet to analyze biofilms formation and bacterial quorum sensing mechanisms of regulation at such small-scale level as a surface of unicellular eukaryotes. The second possibility for bacteria to stay in association with a ciliate is to somehow escape or survive digestion after being phagocytized.

It is well known that many pathogenic bacteria enter the host cells by phagocytosis Some opportunistic bacteria, if engulfed by a phagocyte, are also able to avoid digestion. For some bacteria, intracellular persistence may even trigger and increase their pathogenic properties and virulence Thus, protists may serve as transient reservoirs for many potentially dangerous bacteria. At the same time, the similar role of ciliates has been also shown, but small-scale investigations were focused mainly on pathogens found as endosymbionts of ciliates 15 , In some recent works 49 , 61 diverse digestion-resistant bacteria were described in different ciliates; some of them were enough numerous to be visualized inside the host cells with fluorescence in situ hybridization.

However, the abundance of different bacteria forming the microbiome of ciliates is usually low, thus making NGS analysis a much more sensitive technique to study these phenomena. The ciliate core microbiomes then would be mostly formed by such bacteria, and the ratio between these would be dynamic and varying from cell to cell; sometimes also occasional bacteria could dominate, forming a transitory part of a microbiome.

Actually this is what we obtained as a result of our metabarcoding analysis. Thus, ciliates can play in nature the role of reservoirs for potentially opportunistic and pathogenic bacteria. Statistical analyses showed that sets of OTUs from the environmental controls were significantly different from those associated to stentors Figs 5 , 6.

Indeed, statistical analyses, nmMDS and PCoA strongly confirmed that prokaryotic communities of Stentor cells and corresponding environments were clearly separated from each other, being very distant Figs 5 , 6.

However, ciliate cells did not display a homogenous microbiome among themselves. When reads of abundant symbionts were removed from the analyses, it became clear that the microbiomes of single cells had some individual differences. At the same time, our data are insufficient to generalize if all specimens of the same ciliate species isolated from one locality possess similar or diverse microbiomes, but allow to suggest that there is a core microbiome for ciliates of the same species, at least isolated from the same locality.

Further studies using several species of ciliates from the same location and from different origins should be performed to clarify this aspect. However, we suppose that environment does not shape the core microbiomes of ciliates, as the latter in the end have almost nothing in common with the environmental prokaryotic community. Three samples were collected from a sewage stream located in Peterhof, St. Petersburg, Russia Stentor cells were observed in vivo , and identified as S.

Three ciliate specimens were isolated as replicates for each sampling point with a sterile micropipette, washed thoroughly through several passages in sterile Volvic water and then kept in the last water aliquot overnight to reduce the load of random bacteria present in food vacuoles. Then stentors were washed briefly in autoclaved sterile distilled water, in order to reduce contaminants and microorganisms attached to the cell surface. Two of nine Stentor samples did not yield sufficient DNA for library preparation and were discarded.

De-multiplexing and quality filtering were performed removing any low quality or ambiguous reads, and sequences shorter than nucleotides were discarded. The most common sequence was selected in each OTU as representative. QIIME was used to evaluate diversity of the prokaryotic communities both within and between samples. Firstly, the relative abundances of prokaryotic taxa were estimated as the percentage of contigs number for each taxon for the environmental and the Stentor cell samples, and the community richness was evaluated calculating the Shannon index.

The significance of OTUs presence in the ciliates samples was assessed using the non-parametric Kruskal-Wallis test. In addition, beta diversity was estimated to compare the bacterial communities of the Stentor cells and their environments. To confirm the significance of differences observed between bacterial communities of the environments and stentors, ANOSIM was calculated.

The same analyses were carried out also after excluding the most abundant OTUs representing symbiotic microorganisms associated with stentors see Results.

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Do ruminal ciliates select their preys and prokaryotic symbionts? Up to 2 mm long, they are larger than some of the smallest multicellular organisms. Stentor usually attach to substrates and form a trumpet shape. If they are free-swimming, they assume an oval or pear shape.

A major characteristic of Stentor is the rapid contraction and extension of the cell body. Cilia mainly cover the rim of the trumpet. In this way, Stentor resemble Vorticella. However, Stentor also have horizontal rows of cilia on the rest of the body to allow it to swim. Contraction and extension are controlled by two longitudinal fiber systems that are found in the cell cortex.

They are called km fibers and myonemes. The primary photosensor is called stentorin. The macronucleus is an important feature of Stentor. It resembles a string of beads, consisting of nodules held together by cytoplasmic bridges. Stentor are omnivorous heterotrophs.

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