ciliates

繊毛虫類、繊毛虫、線毛虫類、線毛虫

関

Ciliophora

WordNet

1. having a margin or fringe of hairlike projections (同)ciliated
2. a protozoan with a microscopic appendage extending from the surface of the cell (同)ciliated protozoan, ciliophoran

PrepTutorEJDIC

1. 繊毛虫(ゾウリムシなど,繊毛のある微生物)

Wikipedia preview

出典(authority):フリー百科事典『ウィキペディア（Wikipedia）』「2015/05/23 00:42:35」(JST)

wiki en

[Wiki en表示]

This article is about the protozoan phylum Ciliophora. For the type of ciliate cells in general, see Ciliate cells. For the type of leaf margin, see Leaf#Margins (edge).

Ciliates Temporal range: Ediacaran–Recent PreꞒ Ꞓ O S D C P T J K Pg N
"Ciliata" from Ernst Haeckel's Kunstformen der Natur, 1904
Scientific classification
Domain:	Eukarya
Kingdom:	Chromalveolata
Superphylum:	Alveolata
Phylum:	Ciliophora Doflein, 1901 emend.
Classes
Karyorelictea Heterotrichea Spirotrichea Litostomatea Phyrea Plagiopylea See text for subclasses.
Synonyms
Ciliata Perty, 1852

The ciliates are a group of protozoans characterized by the presence of hair-like organelles called cilia, which are identical in structure to eukaryotic flagella, but in general shorter and present in much larger numbers, with a different undulating pattern than flagella. Cilia occur in all members of the group (although the peculiar Suctoria only have them for part of the life-cycle) and are variously used in swimming, crawling, attachment, feeding, and sensation. Nevertheless, although not a general practice, some authors prefer to call "cilia" both eukaryotic cilia and flagella, reserving the term "flagella" to the bacterial structure. Anywise, the fundamental difference between multiciliate flagellates (e.g., hemimastigids, Stephanopogon, Multicilia, opalines) and ciliates is the presence of macronuclei in ciliates alone.

Ciliates are an important group of protists, common almost everywhere there is water — in lakes, ponds, oceans, rivers, and soils. About 3,500 species have been described, and the potential number of extant species is estimated at 30,000. Included in this number are many ectosymbiotic and endosymbiotic species, as well as some obligate and opportunistic parasites. Ciliate species range in size from as little as 10 µm to as much as 4 mm in length, and include some of the most morphologically complex protozoans.

In most systems of taxonomy, "Ciliophora" is ranked as a phylum, under either the kingdom Protista or Protozoa. In some systems of classification, ciliated protozoa are placed within the class "Ciliata," (a term which can also refer to an unrelated genus of fish). In the taxonomic scheme proposed by the International Society of Protistologists, which eliminates formal rank designations such as "phylum" and "class", "Ciliophora" is an unranked taxon within Alveolata.

Cell structure

Cell structure of a ciliophoran: 1-contractile vacuole, 2-digestive vacuole, 3-macronucleus, 4-micronucleus, 5-cytoproct, 6-cytopharynx, 7-cytostome, 8-cilium.

Nuclei

Unlike most other eukaryotes, ciliates have two different sorts of nuclei: a small, diploid micronucleus (reproduction), and a large, polyploid macronucleus (general cell regulation). The latter is generated from the micronucleus by amplification of the genome and heavy editing. The micronucleus serves as the germ line nucleus, but does not express its genes. The macronucleus provides the nuclear RNA for vegetative growth.

Division of the macronucleus occurs by amitosis and the segregation of the chromosomes occurs by a process whose mechanism is unknown. This process is by no means perfect, and after about 200 generations the cell shows signs of aging. Periodically the macronuclei must be regenerated from the micronuclei. In most, this occurs during conjugation. Here two cells line up, the micronuclei undergo meiosis, some of the haploid daughters are exchanged and then fuse to form new micronuclei and macronuclei.

Cytoplasm

Food vacuoles are formed through phagocytosis and typically follow a particular path through the cell as their contents are digested and broken down by lysosomes so the substances the vacuole contains are then small enough to diffuse through the membrane of the food vacuole into the cell. Anything left in the food vacuole by the time it reaches the cytoproct (anus) is discharged by exocytosis. Most ciliates also have one or more prominent contractile vacuoles, which collect water and expel it from the cell to maintain osmotic pressure, or in some function to maintain ionic balance. In some genera, such as Paramecium, these have a distinctive star shape, with each point being a collecting tube.

Specialized structures in ciliates

In some forms there are also body polykinetids, for instance, among the spirotrichs where they generally form bristles called cirri. More often body cilia are arranged in mono- and dikinetids, which respectively include one and two kinetosomes (basal bodies), each of which may support a cilium. These are arranged into rows called kineties, which run from the anterior to posterior of the cell. The body and oral kinetids make up the infraciliature, an organization unique to the ciliates and important in their classification, and include various fibrils and microtubules involved in coordinating the cilia.

The infraciliature is one of the main component of the cell cortex. Another are the alveoli, small vesicles under the cell membrane that are packed against it to form a pellicle maintaining the cell's shape, which varies from flexible and contractile to rigid. Numerous mitochondria and extrusomes are also generally present. The presence of alveoli, the structure of the cilia, the form of mitosis and various other details indicate a close relationship between the ciliates, Apicomplexa, and dinoflagellates. These superficially dissimilar groups make up the alveolates.

Feeding

Most ciliates are heterotrophs, feeding on smaller organisms, such as bacteria and algae, and detritus swept into the oral groove (mouth) by modified oral cilia. This usually includes a series of membranelles to the left of the mouth and a paroral membrane to its right, both of which arise from polykinetids, groups of many cilia together with associated structures. The food is moved by the cilia through the mouth pore into the gullet, which forms food vacuoles.

This varies considerably, however. Some ciliates are mouthless and feed by absorption (osmotrophy), while others are predatory and feed on other protozoa and in particular on other ciliates. Some ciliates parasitize animals, although only one species, Balantidium coli, is known to cause disease in humans.

Reproduction and sexual phenomena

Ciliate undergoing the last processes of binary fission, a form of asexual reproduction

Ciliates reproduce asexually, by various kinds of fission. During fission, the micronucleus undergoes mitosis and the macronucleus elongates and splits in half (except among the Karyorelictean ciliates, whose macronuclei do not divide). The cell then divides in two, and each new cell obtains a copy of the micronucleus and the macronucleus.

Typically, the cell is divided transversally, with the anterior half of the ciliate (the proter) forming one new organism, and the posterior half (the opisthe) forming another. However, other types of fission occur in some ciliate groups. These include budding (the emergence of small ciliated offspring, or "swarmers," from the body of a mature parent); strobilation (multiple divisions along the cell body, producing a chain of new organisms); and palintomy (multiple fissions, usually within a cyst).

Fission may occur spontaneously, as part of the vegetative cell cycle. Alternatively, it may be proceed as a result of self-fertilization (autogamy), or it may follow conjugation, a sexual phenomenon in which ciliates of compatible mating types exchange genetic material.

Conjugation

Overview

During conjugation, two cells form a bridge between their cytoplasms, the micronuclei undergo meiosis, the macronuclei disappear, and the haploid micronuclei are exchanged over the bridge. In some ciliates (such as Vorticella), conjugating cells become permanently fused, and one conjugant is absorbed by the other. In most ciliate groups, however, the cells separate after conjugation, and both form new macronuclei from their micronuclei. Conjugation and autogamy are always followed by fission.

More details

In general the process is as follows but the details may vary between species:

• Compatible mating strains meet and partly fuse.
• The micronuclei undergo meiosis producing four micronuclei per cell.
• Three of these micronuclei disintegrate. The fourth undergoes mitosis.
• The two cells exchange a micronucleus.
• The cells then separate.
• The micronuclei in each cell fuse.
• This is followed by mitosis which occurs three times giving rise to eight micronuclei.
• Four of the new micronuclei transform into macronuclei
• Finally binary fission occurs twice yielding four daughter cells.

DNA Rearrangements (gene scrambling)

Development of the Oxytricha macronuclear genome from micronuclear genome

Ciliates contain two types of nuclei: somatic "macronucleus" and the germline "micronucleus". Only the DNA in the micronucleus is passed on during sexual reproduction (conjugation). On the other hand, only the DNA in the macronucleus is actively expressed and results in the phenotype of the organism. Macronuclear DNA is derived from micronuclear DNA by amazingly extensive DNA rearrangement and amplification.

The macronucleus begins as a copy of the micronucleus. The micronuclear chromosomes are fragmented into many smaller pieces and amplified to give many copies. The resulting macronuclear chromosomes often contain only a single gene. In Tetrahymena, the micronucleus has 10 chromosomes (5 per haploid genome), while the macronucleus has over 20,000 chromosomes.

In addition, the micronuclear genes are interrupted by numerous "Internal Eliminated Sequences" (IESs). During development of the macronucleus, IESs are deleted and the remaining gene segments, Macronuclear Destined Sequences (MDSs), are spliced together to give the operational gene. Tetrahymena has about 6000 IESs and about 15% of micronuclear DNA is eliminated during this process. The process is guided by small RNAs and epigenetic chromatin marks.

In spirotrich ciliates (such as Oxytricha), the process is even more complex due to "gene scrambling": the MDSs in the micronucleus are often in different order and orientation from that in the macronuclear gene, and so in addition to deletion, DNA inversion and translocation are required for "unscrambling". This process is guided by long RNAs derived from the parental macronucleus. More than 95% of micronuclear DNA is eliminated during spirotrich macronuclear development.

Fossil record

Until recently, the oldest ciliate fossils known were tintinnids from the Ordovician Period. In 2007, Li et al. published a description of fossil ciliates from the Doushantuo Formation, about 580 million years ago, in the Ediacaran Period. These included two types of tintinnids and a possible ancestral suctorian.

Classification

Stentor roeseli

Subphylum Postciliodesmatophora

• Class Karyorelictea
• Class Heterotrichea (e.g. Stentor)

Subphylum Intramacronucleata

Oxytricha trifallax

• Class Spirotrichea
  • Subclass Choreotrichia (e.g. Tintinnidium)
  • Subclass Oligotrichia (e.g. Halteria)
  • Subclass Stichotrichia (e.g. Stylonychia)
  • Subclass Hypotrichia (e.g. Euplotes)

A trophozoite of Balantidium coli

• Class Litostomatea
  • Subclass Haptoria (e.g. Didinium)
  • Subclass Trichostomatia (e.g. Balantidium)
• Class Phyllopharyngea
  • Subclass Phyllopharyngia (e.g. Chilodonella uncinata
  • Subclass Rhynchodia
  • Subclass Chonotrichia
  • Subclass Suctoria (e.g. Podophrya)
• Class Nassophorea
• Class Colpodea (e.g. Colpoda)
• Class Prostomatea (e.g. Coleps)
• Class Oligohymenophorea
  • Subclass Peniculia (e.g. Paramecium)
  • Subclass Hymenostomatia (e.g. Tetrahymena)
  • Subclass Scuticociliatia
  • Subclass Peritrichia (e.g. Vorticella)
  • Subclass Astromatia
  • Subclass Apostomatia
• Class Plagiopylea

Other

• Class Prostomatea
  • Genus Cryptocaryon

Some old classifications included Opalinidae in the ciliates.

References

• "Ciliophora". NCBI Taxonomy Browser. 5878. 

UpToDate Contents

全文を閲覧するには購読必要です。 To read the full text you will need to subscribe.

• 1. 抗原虫治療 antiprotozoal therapies

English Journal

• Variability in Secondary Structure of 18S Ribosomal RNA as Topological Marker for Identification of Paramecium species.

• Shakoori FR1, Tasneem F, Al-Ghanim K, Mahboob S, Al-Misned F, Jahan N, Shakoori AR.
• Journal of cellular biochemistry.J Cell Biochem.2014 Dec;115(12):2077-88. doi: 10.1002/jcb.24885.
• Besides cytological and molecular applications, Paramecium is being used in water quality assessment and for determination of saprobic levels. An unambiguous identification of these unicellular eukaryotes is not only essential, but its ecological diversity must also be explored in the local environm
• PMID 25043709

• The Food Web of Boiling Springs Lake Appears Dominated by the Heterolobosean Tetramitus thermacidophilus strain BSL.

• Reeder WH 4th1, Sanck J, Hirst M, Dawson SC, Wolfe GV.
• The Journal of eukaryotic microbiology.J Eukaryot Microbiol.2014 Nov 7. doi: 10.1111/jeu.12193. [Epub ahead of print]
• We studied the protist grazers of Boiling Springs Lake (BSL), an acid geothermal feature in Lassen Volcanic National Park, using a combination of culture and genetic approaches. The major predator in BSL is a vahlkampfiid amoeba closely related (95% 18S+ITS rRNA identity) to Tetramitus thermacidophi
• PMID 25382699

• Morphology, Ontogeny, and Phylogeny of Two Brackish Urostylid Ciliates (Protist, Ciliophora, Hypotricha).

• Chen X1, Miao M, Ma H, Al-Rasheid KA, Xu K, Lin X.
• The Journal of eukaryotic microbiology.J Eukaryot Microbiol.2014 Nov;61(6):594-610. doi: 10.1111/jeu.12137. Epub 2014 Aug 25.
• The diversity of hypotrichous ciliates has encouraged numerous researchers to use a combination of morphological, morphogenetic, and phylogenetic data to provide a better understanding of the evolutionary relationships within this complex group. In this study, we investigate the morphology and morph
• PMID 25040093

Japanese Journal

• ASIA/OCEANIA REPORT : Compositional differences among planktonic ciliate communities in four subtropical eutrophic lakes in China

• Li Jing,Chen Feizhou,Liu Zhengwen [他]
• Limnology 14(1), 105-116, 2013-01-00
• NAID 40019552428

• 繊毛虫の生命現象を司る鍵物質

• 粳間 由幸,世良 佳彦,臼杵 克之助,飯尾 英夫
• 有機合成化学協会誌 71(3), 207-216, 2013
• … Protozoan ciliates are eukaryotic unicellular organisms characterized by the presence of hair-like organelles called cilia. …
• NAID 130003373272

• Revealing the Diversity and Quantity of Peritrich Ciliates in Environmental Samples Using Specific Primer-based PCR and Quantitative PCR

• LIU XIHAN,GONG JUN
• Microbes and environments 27(4), 497-503, 2012-12-01
• … The development of peritrich-specific primers, for the first time, for conventional PCR and qPCR assays, provides useful molecular tools for revealing the diversity and quantity of peritrich ciliates in environmental samples. …
• NAID 10031125950

Related Links

• ciliate | protozoan | Britannica.com

Any member of the protozoan phylum Ciliophora, of which there are some 8,000 species; ciliates are generally considered the most evolved and complex of protozoans. Ciliates are... ... We welcome suggested improvements to any ...

• Ciliates - Microscopy-UK Home (Resources for the ...

This movie shows the basic body plan of the ciliate Paramecium. It has cilia all over its cell. What are Ciliates? Ciliates are unicellular protists that can be recognised by their hairlike 'cilia'. They use them for locomotion and for feeding.

Related Pictures

---

★リンクテーブル★

リンク元	「繊毛虫」「繊毛虫類」「線毛虫」「線毛虫類」「Ciliophora」
関連記事	「ciliate」

「繊毛虫」

　　[★]

英

ciliates、Ciliophora

関

線毛虫類、線毛虫、繊毛虫類、繊毛虫門

「繊毛虫類」

　　[★]

英

ciliates

ラ

ciliata

関

線毛虫類、線毛虫、繊毛虫

「線毛虫」

　　[★]

英

ciliates

関

線毛虫類、繊毛虫、繊毛虫類

「線毛虫類」

　　[★]

英

ciliates

関

線毛虫、繊毛虫、繊毛虫類

「Ciliophora」

　　[★]

• 繊毛虫、繊毛虫門

関

ciliates

「ciliate」

　　[★]

• 繊毛性の、有繊毛の

関

ciliated