Holotricha is an order of ciliates. The classification has fallen from use as a formal taxon, but the terms "holotrich" and "holotrichous" are still applied descriptively to organisms with cilia of uniform length distributed evenly over the surface of the body. [1]
The term holotrich derives from the ancient greek ὅλος (hólos), meaning "whole, entire", and θρίξ , τριχός (thríx, trikhós), meaning 'hair', [2] [3] because of the even distribution of cilia over the surface of the cell. [1]
The order Holotricha was erected in 1859 by Friedrich Stein, who developed a system of classification in which ciliates were categorized according to the type and distribution of their cilia. The four major subdivisions of Stein's Ciliata were: Heterotricha, Peritricha, Hypotricha and Holotricha. The latter was reserved for organisms with cilia covering the entire surface of the body. [4] The order included five families, within which a very diverse collection of genera were grouped. Among the better-known ciliates in Stein's Holotricha were Coleps , Nassula , Lacrymaria , Loxodes , Colpoda , Glaucoma and Paramecium . [5]
Stein's scheme was amended in 1889 by Otto Bütschli, who divided the Ciliata into two orders: Holotricha and Spirotricha. Bütschli's Holotricha were described much as Stein's had been, as ciliates with relatively uniform somatic cilia. The other three orders of Stein's Ciliata were placed in a lower rank under the new order Spirotricha: ciliates possessing a well-developed structure of spiralling membranelles around the oral cavity. [6]
Versions of Bütschli's two-order scheme were used by taxonomists throughout the first half of the twentieth century, although many adjustments were made to the arrangement of families and genera within those groups. [7] [8] [9] Meanwhile, improved laboratory methods and the advent of the electron microscope began to reveal inadequacies in the old scheme; and, as inconsistencies accumulated, dissatisfaction with the system grew. [10] Nonetheless, Holotricha remained in use as a high-level taxon until 1974, when John Corliss proposed a complete revision of the phylum Ciliophora. In his system, Spirotricha is retained as an order, under the class Polyhymenophora, but Holotricha does not appear at any rank. Recent systems of classification, including an influential scheme proposed by Pierre de Puytorac in 1994, and Denis Lynn's revision of 2002, make no mention of Holotricha. [11]
The spirotrichs are a large and diverse group of ciliate protozoa. They typically have prominent oral cilia in the form of a series of polykinetids, called the adoral zone of membranelles, beginning anterior to the oral cavity and running down to the left side of the mouth. There may also be one or two paroral membranes on its right side. The body cilia are fused to form polykinetids called cirri in some, and are sparse to absent in others.
The alveolates are a group of protists, considered a major clade and superphylum within Eukarya. They are currently grouped with the stramenopiles and Rhizaria among the protists with tubulocristate mitochondria, the group being referred to as SAR.
The heterotrichs are a class of ciliates. They typically have a prominent adoral zone of membranelles circling the mouth, used in locomotion and feeding, and shorter cilia on the rest of the body. Many species are highly contractile, and are typically compressed or conical in form. These include some of the largest protozoa, such as Stentor and Spirostomum, as well as many brightly pigmented forms, such as certain Blepharisma.
The plagiopylids are a small order of ciliates, including a few forms common in anaerobic habitats.
The Oligohymenophorea are a large class of ciliates. There is typically a ventral groove containing the mouth and distinct oral cilia, separate from those of the body. These include a paroral membrane to the right of the mouth and membranelles, usually three in number, to its left. The cytopharynx is inconspicuous and never forms the complex cyrtos found in similar classes. Body cilia generally arise from monokinetids, with dikinetids occurring in limited distribution over part of the body.
The Phyllopharyngea are a class of ciliates, including some which are extremely specialized. Motile cells typically have cilia restricted to the ventral surface, or some part thereof, arising from monokinetids with a characteristic ultrastructure. In both chonotrichs and suctoria, however, only newly formed cells are motile and the sessile adults have undergone considerable modifications of form and appearance. Chonotrichs, found mainly on crustaceans, are vase-shaped, with cilia restricted to a funnel leading down into the mouth. Mature suctorians lack cilia altogether, and initially were not classified as ciliates.
The hypotrichs are a group of ciliated protozoa, common in fresh water, salt water, soil and moss. Hypotrichs possess compound ciliary organelles called "cirri," which are made up of thick tufts of cilia, sparsely distributed on the ventral surface of the cell. The multiple fused cilia which form a cirrus function together as a unit, enabling the organism to crawl along solid substrates such as submerged debris or sediments. Hypotrichs typically possess a large oral aperture, bordered on one side by a wreath or collar of membranelles, forming an "adoral zone of membranelles," or AZM.
The stichotrichs were a proposed group of ciliates, in the class Spirotrichea. In a classification system proposed by Eugene Small and Denis Lynn in 1985, Stichotrichia formed a subclass containing four orders: Stichotrichida, Urostylida, Sporadotrichida and Plagiotomida. Although the group was made up of species traditionally classified among the "hypotrichs"--ciliates possessing compound ciliary organelles called cirri--it excluded euplotid ciliates such as Euplotes and Diophrys, which were placed in the subclass Hypotrichia. In later classifications proposed by Denis Lynn, Stichotrichia omits the order Plagiotomida.
Membranelles are structures found around the mouth, or cytostome, in ciliates. They are typically arranged in series, to form an "adoral zone of membranelles," or AZM, on the left side of the buccal cavity (peristome). The membranelles are made up of kinetosomes arranged in groups to make up polykinetids. The cilia which emerge from these structures appear to be fused and to function as a single membrane, which can be used to sweep particles of food into the cytostome, or for locomotion.
Didinium is a genus of unicellular ciliates with at least ten accepted species. All are free-living carnivores. Most are found in fresh and brackish water, but three marine species are known. Their diet consists largely of Paramecium, although they will also attack and consume other ciliates. Some species, such as D. gargantua, also feed on non-ciliate protists, including dinoflagellates, cryptomonads, and green algae.
Protozoa is an informal term for a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Historically, protozoans were regarded as "one-celled animals", because they often possess animal-like behaviours, such as motility and predation, and lack a cell wall, as found in plants and many algae.
Colpoda is a genus of ciliates in the class Colpodea, order Colpodida, and family Colpodidae.
Karyorelictea is a class of ciliates in the subphylum Postciliodesmatophora. Most species are members of the microbenthos community, that is, microscopic organisms found in the marine interstitial habitat, though one genus, Loxodes, is found in freshwater.
The ciliates are a group of alveolates characterized by the presence of hair-like organelles called cilia, which are identical in structure to eukaryotic flagella, but are in general shorter and present in much larger numbers, with a different undulating pattern than flagella. Cilia occur in all members of the group and are variously used in swimming, crawling, attachment, feeding, and sensation.
Euplotes is a genus of ciliates in the subclass Euplotia. Species are widely distributed in marine and freshwater environments, as well as soil and moss. Most members of the genus are free-living, but two species have been recorded as commensal organisms in the digestive tracts of sea urchins.
Cothurnia is a genus of freshwater and marine peritrichs in the family Vaginicolidae. It is characterised by living in a transparent tubular lorica. During the feeding or vegetative phase of its life cycle, Cothurnia attaches to submerged surfaces through a short stalk — mostly on the surfaces of fishes, crustaceans and aquatic plants. It is commonly studied for its epibiotic relationship with the host that it is attached to.
Protoheterotrichida is an order of karyorelict ciliates. It contains the family Geleiidae.
Intramacronucleata is a subphylum of ciliates. The group is characterized by the manner in which division of the macronucleus is accomplished during binary fission of the cell. In ciliates of this subphylum, division of the macronucleus is achieved by the action of microtubules which are assembled inside the macronucleus itself. This is in contrast to heterotrich ciliates of the subphylum Postciliodesmatophora, in which division of the macronucleus relies on microtubules formed outside the macronuclear envelope.
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.
Isotrichidae is a family of single-celled organisms (protozoa) in the order Vestibuliferida. Most species in this family are endosymbionts of ruminant animals. They are holotrichous, that is, their surface is covered in cilia of uniform length, and they contain several contractile vacuoles.