The Fine Structure of the Internal Spiral of Kinetosomes in Foettingeria's Tomite,a Unique Storage Depot for Kinetosomes1

ABSTRACT. The fine structure of the tomite of Foettingeria actiniarum (Claparède) was examined and compared with that of other apostome tomites. This stage in the life cycle has a unique configuration of kineties that form a spiral through the cytoplasm in the interior of the body. The structure and behavior of this internal spiral were evaluated as a mechanism for the storage of kinetosomes, an adaptation to the ciliate's two-host life cycle. The spiral is composed of nine ribbons of laterally compressed kinetosomes that are in contact with a thin electron-dense fibril. Paralleling the kineties of the spiral are conspicuous, swollen lamellae of the rough endoplasmic reticulum; these lamellae contain moderately electron-dense material. The spiral is associated with the large contractile vacuole and winds about the macronucleus. The tomite of Foettingeria possesses a single, robust, caudal cilium located in a pit, along with the nozzle-like pore of the contractile vacuole. The walls of the pit contain several trichocysts arranged radially about the caudal cilium and aimed into the pit.     

Evidence for Hymenostome Affinities in an Apostome Ciliate

The functional mouth of exuviotrophic apostome ciltates appears only after an elaborate metamorphosis that begins at the onset of the molting of their crustacean hosts. In the tomite. a non-feeding migratory stage, a mid-ventrai depression at the origin of kineties x, y and z has been misidentified as the cytostome. Studies of fine structure and morphogenesis identify the true but nonfunctional cytostome—the subapiral lateral canal —and the falciform and ogival fields as the adoral ciliature. The anterior row of barren kinetosomes that parallels on the right the anterior third of the lateral canal is actually the infraciliature of a paroral. 2 rows of barren staggered kinetosomes. The canal itself is a narrow tube, its walls partially lined with microtubules. It begins 2–3 μm from the apex of the body and passes between falciform field 9 and the ogival field to end near the end of the ogival field. The fine structure of the infraciliature of the falciform and ogival fields differs markedly from that of the somatic kineties. In the host's early pre-molt stages, the paroral migrates across the ventral surface of the encysted phoront and is accompanied by the microtubules of the lateral canal. The anterior end of falciform field 9 disorganizes into scattered kinetosomes, the trophont's anterior field of kinetosomes, but the posterior end migrates in an arc across the anterior ventral surface and remains as kinety a located near the angle where kinety 1 sharply par ra continues posteriad ind dorsad to the posterior limit of the extended cytostome. At the end of metamorphosis it sinks into The cytoplasm and disappears. The completion of the extended cytostome, the functional mouth, marks the termination of the microstome-macrostome transformation. The fine structure of the infraciliature and microtubular elements making up the macrostome and the evocation of the microstome-macrostome transformation in the presence of specific foods suggest that apostome ciliates any more properly be a suborder of Hymenostomatida rather than a subclass of Oligohymenophorea.     

Morphologie,Infraciliatur und Ökologie der limnischen Tintinnina: Tintinnidium fluviatile Stein,Tintinnidium pusillum Entz,Tintinnopsis cylindrata Daday und Codonella cratera (Leidy) (Ciliophora,Polyhymenophora)*

Synopsis. Structure, infraciliature, and ecology of 4 fresh-water Tintinnina were investigated. The lorica of Tintinnidium fluviatile is gelatinous, fragile, and contains some agglutinated material mainly of biologic origin. Its infraciliature consists of ? 10 kineties. with kinetosomes arranged in pairs. Only one basal body of a pair is ciliated, except for the uppermost 1–4 pairs which have 2 slightly elongated cilia. In Tintinnidium fluviatile. Tintinnidium pusillum, and Tintinnopsis cylindrata there are 2 prominent ventral organelles. The lorica of T. pusillum is gelatinous and coated with much agglutinated material of biologic and nonbiologic origin. Its infraciliature is similar to that of T. fluviatile, but the uppermost pair of kinetosomes has elongated cilia. The firm loricae of T. cylindrata and Codonella cratera are, built mainly of sharp-cornered structures. The infraciliature of T. cylindrata is composed of ? 10 kineties with kinetosomes not arranged in pairs. The infraciliature of C. cratera consists of ? 32 kineties, in some of which the kinetosomes are paired, e.g. ventral kinety, and in others not paired, e.g. cilia of the very prominent lateral field and of the other somatic kineties. The uppermost kinetosomes of each somatic kinety are paired and have elongated cilia. In addition, there is an unusual ventro-lateral kinety. The oral apparatus consists of adoral membranelles and a paroral membrane. The membranelles that enter the praecral cavity are very elongate, a feature perhaps unique to Polyhymenophora. The fibrillar system consists of a prominent praeoral ring formed by fibrils extending from the adoral membranelles. A finely meshed silverline system extends over the entire cell. A review of the ecology of the fresh-water Tintinnina indicated that water temperature seems to be the most essential ecologic factor. The systematic position of the Tintinnina is discussed in light of their infraciliature. It is concluded that these organisms are most closely related to Oligotrichina, and probably to Heterotrichina.     

The fine structure of secretion in Hyalophysa chattoni: formation of the attachment peduncle and the chitinous phoretic cyst wall.

The settling tomite stage of the apostome Hyalophysa chattoni secretes a phoretic cyst wall composed of chitin, mucopolysaccharides, and protein. Within 1 1/2 h after settling, an electron-dense proteinaceous cyst layer (the outer layer) is formed from secretions originating at the base of the kineties and from the thick pellicular layer between the kineties. The inner cyst layer, composed primarily of chitin (acidic and neutral polysaccharides are also present), is secreted across the entire cell surface. Cyst wall formation is completed within 6 h. The fine structure of endocyst secretion resembles stages in the secretion of chitin by fungi, yeasts, and arthropods. A proteinaceous attachment peduncle is secreted to anchor the cell to a shrimp host and is formed by the release of electrondense dense secretory bodies from the cell's ventral surface.     

Metamorphosis from the Phoront to the Trophont in Hyalophysa*

SYNOPSIS. The nature and sequence of changes involved in the metamorphosis of the phoront of Hyalophysa have been examined in the light microscope using both living and silver-impregnated ciliates. The phoront's infraciliature, which resembles that of the migratory tomite, differentiates to the trophont (feeding stage) infraciliature without an intervening dedifferentiation. The primary visible event of metamorphosis is the growth of the metastomial area that distorts the meridional somatic kineties so that they curve or spiral around it. The distinct borders and intense argentophilia of the metastomial area suggest that it is a discrete organelle which is probably involved in the management and concentration of a large volume of dilute food. The anterior ventral field of kinetosomes, whose presence distinguishes Hyalophysa from Gymnodinioides, forms from a disorganized anterior segment of falciform field 9. The rapid and pronounced elongation of the somatic kineties supports our view that the kinetodesmos elongates by the sliding of its component subfibrils upon one another, and that the accessory kinetosomes seen in the electron microscope become functional at metamorphosis. The movement of the kineties and contractile vacuole pore during metamorphosis suggests that differentiation in this instance is not directed by pellicular “fields” but by morphogenetic movements of the underlying cytoplasm.     

The Fine Structure of Secretion in Hyalophysa chattoni: Formation of the Attachment Peduncle and the Chitinous Phoretic Cyst Wall

ABSTRACT. The settling tomite stage of the apostome Hyalophysa chattoni secretes a phoretic cyst wall composed of chitin, mucopolysaccharides, and protein. Within 1 1/2 h after settling, an electron-dense proteinaceous cyst layer (the outer layer) is formed from secretions originating at the base of the kineties and from the thick pellicular layer between the kineties. The inner cyst layer, composed primarily of chitin (acidic and neutral polysaccharides are also present), is secreted across the entire cell surface. Cyst wall formation is completed within 6 h. The fine structure of endocyst secretion resembles stages in the secretion of chitin by fungi, yeasts, and arthropods. A proteinaceous attachment peduncle is secreted to anchor the cell to a shrimp host and is formed by the release of electron-dense secretory bodies from the cell's ventral surface.     

Cortical and Nuclear Events During Cell Division and Resting Cyst Formation in Colpoda inflata

Nuclear and cortical phenomena during dividing and resting cyst formation of Colpoda inflata are described. Cell division forms a cyst and produces two or four tomites. In each tomite, the right oral field results from the proliferation of the anterior extreme of a single kinety, and the left oral field results from the proliferation of four, five, or six somatic kineties. After macronuclear division, each macronuclear mass undergoes a chromatinic extrusion process. During resting cyst formation, the oral infraciliature of the vegetative cell is resorbed. The somatic kineties dispose in a radial way and some pairs of kinetosomes disappear. As in cell division, there is an extrusion process. From these results we conclude that the resting cysts of Colpoda inflata cannot be included in any group of the previous classifications for hypotrich resting cysts. Thus, we propose a new additional group to Walker and Maugel's classification called PKR (partial-kinetosome-resorbing) cysts.     

The Life Cycle and Morphology of the Apostomatous Ciliate,Hyalophysa chattoni n. g., n. sp.*

SYNOPSIS. A new apostome ciliate was discovered in collections at Friday Harbor, Washington and the San Francisco area. All stages of the life cycle were studied in both living and stained condition. Dormant encysted stages (phoronts) occur on the gills of Pagurus hirsutiusculus. Excystation occurs in synchrony with the molting of the host yielding the trophic stage (trophont), which feeds on the exuvial fluids trapped in the crab's cast-off exoskeleton. The trophont becomes greatly enlarged as a result of feeding, and the cytoplasm and organelles become compressed into a thin cortical layer. Each fully grown trophont encysts (becoming a tomont) as a prelude to repeated binary fission, which results in the release of actively motile offspring (tomites). These disperse and promptly resume the encysted phoront stage on the host's gills. The Chatton-Lwoff silver impregnation method revealed that all stages of the life cycle have nine somatic kineties. In the trophont stage they are accompanied by an anterior ventral field of scattered clumps of kinetosomes. During conjugation the partners attach by their ventral surfaces between kineties 1 and 9 and at the left of the ventral field. The tomite stage was stained with Protargol. In addition to the characteristic features of the foettingeriid tomite also revealed by the Chatton-Lwoff method, Protargol revealed the following heretofore undescribed morphological features: a short row of kinetosomes immediately anterior to the ogival field; a line paralleling the left margin of the field; the continuity of kinety 8 with falciform field 8; the entrance of kinety 9 into the mouth and its ending against the rosette (an enigmatic organelle characteristic of Foettingeriinae). Feulgen stains showed that the chromatin in the macronucleus is dispersed in aggregates whose size and number vary with the stage of the life cycle. The major period of chromatin synthesis appears to be during the early tomont stage, when Feulgen-positive material increases visibly in amount and intensity of staining. This apostome ciliate was characterized as a new genus on the basis of the infraciliature of the trophont stage, its conjugation with ventral surfaces appressed, and its life cycle. It is named Hyalophysa (hyalo = glassy, physa = bubble) chattoni.     

Oral Regeneration in the Ciliate Stentor coeruleus: A Scanning and Transmission Electron Optical Study

SYNOPSIS. Elaboration of ciliated feeding organelles in the protozoon Stentor coeruleus was reinvestigated for the first time by scanning electron microscopy which gives the most realistic 3-dimensional images. Parallel transmission EM studies of synchronized regenerating stentors gave further ultrastructural details of stomatogenesis, while also confirming the expectation that in the structure of its kineties this now classical experimental object does not differ from other species of Stentor previously studied. Within 2 hr after the stimulus to regeneration, several generations of new kinetosomes for the oral primordium are produced, first in association with kinetosomes of kineties at the restricted primordium site. These kinetosomes rapidly sprout membranellar cilia as well as subpellicular microtubules but are still randomly oriented (anarchic field). The forming membranellar band increases from its center-line to both sides while it grows in length. Young cilia are blunt-ended. Recession of the early anlage occurs without rupture of the pellicle; soon apparent is the clear border stripe of unknown function along the right side of the membranellar band. Instantaneous fixation of beating cilia in early primordia revealed random beating, with coordination and presumably membranellar organization not yet attained. In late anlagen there are 2 types of metachronal rhythm: transversely from cilium to cilium across any given membranelle, as well as the easily observable serial beating of membranelles along the entire band. A single file of cilia leads the subsequent cytostomal invagination. The posterior end of the membranellar band then follows to line the cytopharynx.     

The Fine Structure of the Phoront of the Apostomatous Ciliate,Hyalophysa chattoni*

The phoront of the apostomatous ciliate, Hyalophysa chattoni, is an encysted stage that is carried on the exoskeleton of its crustacean host until the ecdysis of the host. At molting the phoront rapidly metamorphoses to the feeding stage, excysts, and immediately begins to feed on exuvial fluid trapped in the cast-off exoskeleton. The fine structure of the resting phoront resembles that of the preceding migratory stage, the tomite. A prominent ventral tuft of cilia, the ogival field, has vanished, and the trichocysts that paralleled the kinetics have disappeared. The dense inclusion bodies that were concentrated around the mouth and falciform fields have dispersed and greatly decreased in number. The cytoplasm and its membranous organelles do not appear visibly condensed or altered from the preceding stage in the life cycle. The phoront is merely quiescent instead of dormant. Unlike the few ciliate cysts previously examined by electron microscopy, the phoront's cyst is not divisible into separable layers. It resembles the loricae of certain suctoria in being formed principally of a fibrous substance, the outer surface of which has a paracrystalline pattern. The peduncle attaching the cyst to the crab's gill is a continuation of the cyst wall although its structure is somewhat modified. The most conspicuous innovation in the phoront's fine structure is the massive tracts of microtubules that run longitudinally through the macronucleus. The microtubules are in intimate contact with Feulgen-positive chromatin masses which are crowded toward the periphery of the macronucleus.     

Ultrastructure of the cytoplasm of the lower holotrichous infusorian Tracheloraphis prenanti]

The ciliature of T. prenanti Dragesco 1960 (forma oligocineta Raikov et Kovaleva, 1968) consists of 14-18 ventral and lateral longitudinal kineties with paired kinetosomes, carrying either two cilia or one cilium per kinetosome pair (in the latter case, the nonciliated kinetosome is always the posterior one). The ectoplasmic fibrillar system belongs to the postciliary type. A pair of kinetosomes shares a common basal plate. The anterior kinetosome gives rise to a short ribbon of transverse microtubules, the posterior one, to a poorly developed kinetodesmal filament and to a strong ribbon of postciliary microtubules. The latter proceeds backwards along 8 to 12 kinetosome pairs, being incorporated into a laminated postciliodesma which accompanies each kinety on its right side. Rows of Golgi elements, sending secretory vesicles and channels towards the body surface, exist beneath the kinetosome bases. Each kinety is accompanied on its left by a microfibrillar myoneme, surrounded by perimyary vesicles and underlain by a row of mitochondria. The median part of the dorsal surface is nonciliated; the cytoplasm here is rich of membrane systems, contains peripheral, electron-dense, extrusible inclusions and sometimes also bacteria. The electron-dense inclusions develop in the endoplasm, in close contact with mitochondria. The endoplasm contains also large microfibrillar spheres of unknown nature.     

The Kinetid Structures of the Choreotrichous Ciliate Strobilidium velox and an Assessment of Its Evolutionary Lineage1

The ciliary (kinetid) structures of the ciliate Strobilidium velox have been examined with scanning and transmission electron microscopes. Somatic kineties consist of a linear row of kinetosomes (monokinetids) and short cilia lying partially beneath a thin fold of cytoplasm. The only fibrillar kinetid structure extending from the kinetosomes is a transverse ribbon of microtubules. The paroral membrane is a single-file polykinetid possessing a possible transverse ribbon of microtubules and a nematodesma. The oral polykinetids or membranelles are complex, with microtubules extending from both anterior and posterior rows of cilia. While the kinetid structures do not satisfy the criteria for the order Choreotrichida, they are similar to the tintinnids in several other relevant ways. Strobilidium velox is proposed to be an unusual ciliate that is an exception to the concept that somatic kinetids are conservative and reliable phylogenetic indicator structures.     

THE FINE STRUCTURE OF CORTICAL COMPONENTS OF PARAMECIUM MULTIMICRONUCLEATUM

The electron microscope was used to study the structure and three dimensional relationships of the components of the body cortex in thin sections of Paramecium multimicronucleatum. Micrographs of sections show that the cortex is covered externally by two closely apposed membranes (together ~250 A thick) constituting the pellicle. Beneath the pellicle the surface of the animal is molded into ridges that form a polygonal ridgework with depressed centers. It is these ridges that give the surface of the organism its characteristic configuration and correspond to the outer fibrillar system of the light microscope image. The outer ends of the trichocysts with their hood-shaped caps are located in the centers of the anterior and posterior ridges of each polygon. The cilia extend singly from the depressed centers of the surface polygons. Each cilium shows two axial filaments with 9 peripheral and parallel filaments embedded in a matrix and the whole surrouned by a thin ciliary membrane. The 9 peripheral filaments are double and these are evenly spaced in a circle around the central pair. The ciliary membrane is continuous with the outer member of the pellicular membrane, whereas the plasma membrane is continuous with the inner member of the pellicular membrane. At the level of the plasma membrane the proximal end of the cilium is continuous with its tube-shaped basal body or kinetosome. The peripheral filaments of the cilium, together with the material of cortical matrix which tends to condense around them, form the sheath of the basal body. The kinetodesma connecting the ciliary kinetosomes (inner fibrillar system of the light microscopist) is composed of a number of discrete fibrils which overlap in a shingle-like fashion. Each striated kinetosomal fibril originates from a ciliary kinetosome and runs parallel to other kinetosomal fibrils arising from posterior kinetosomes of a particular meridional array. Sections at the level of the ciliary kinetosomes reveal an additional fiber system, the infraciliary lattice system, which is separate and distinct from the kinetodesmal system. This system consists of a fibrous network of irregular polygons and runs roughly parallel to the surface of the animal. Mitochondria have a fine structure similar in general features to that described for a number of mammalian cell types, but different in certain details. The structures corresponding to cristae mitochondriales appear as finger-like projections or microvilli extending into the matrix of the organelle from the inner membrane of the paired mitochondrial membrane. The cortical cytoplasm contains also a particulate component and a system of vesicles respectively comparable to the nucleoprotein particles and to the endoplasmic reticulum described in various metazoan cell types. An accessory kinetosome has been observed in oblique sections of a number of non-dividing specimens slightly removed from the ciliary kinetosome and on the same meridional line as the cilia and trichocysts. Its position corresponds to the location of the kinetosome of the newly formed cilium in animals selected as being in the approaching fission stage of the life cycle.     

Somatic kinetics or paroral membrane: which came first in ciliate evolution?

The ciliate species which lack a distinctive oral ciliature are considered to represent an ancestral state in ciliate evolution. Consequently, the somatic kineties composed of kinetids (kinetosomes plus cilia and associated fibrillar systems) are thought to be the ancestral ciliature. Results on stomatogenesis in 'gymnostomial ciliates' have shown that these ciliates probably have evolved from ancestors already equipped with an oral ciliature. Thus instead of the somatic, the oral ciliature may be regarded an ancestral. Based on these ideas a hypothesis on the evolution of the ciliate kinetome (assembly of all kinetids covering the body of a given ciliate) is presented. The first step in the evolution of the kinetome was the formation of a paroral membrane, a compound ciliary organelle lying along the right side of the oral area which historically but falsely is termed membrane. It was composed of kinetosomal dyads (dikinetids), derived from the kinetid of a dinoflagellate-like ancestor. From the beginning the paroral membrane was responsible for locomotion, ingestion and for the formation of a cytopharyngeal tube which the first ciliate probably had inherited from its flagellate ancestor. In the second step a first somatic kinety was formed from the right row of kinetosomes of the paroral membrane as a result of a longitudinal splitting of the paroral membrane and a subsequent migration of the forming kinety to the right into the somatic cortex. To increase the number of somatic kineties this process was repeated until the kinety produced first reached the left border of the oral area. By this step the locomotive and the nutritional functions were differentiated between somatic and oral structures. In a third step the adoral organelles were formed from somatic kinetids left of the oral area. The primitive type of stomatogenesis was a buccokinetal one derived from the mode the flagellate ancestor used to distribute its replicated kinetosomes to the offspring cells (buccokinetal means that at least parts of the oral anlage for the posterior offspring cell has its origin in the parental oral apparatus). This hypothesis, based on comparative studies on ciliate morphogenesis, is corroborated by molecular data from other laboratories.     

Gymnodinioides pitelkae n. sp. (Ciliophora,Apostomatina) from the littoral amphipod,Marinogammarus obtusatus,a trophont with remnants of the tomite's infraciliature

The apostome genus, Hyalophysa, is a symbiont on most littoral decapods in the US, but no other crustacean orders have been investigated. When exuvia of the amphipod Marinogammarus obtusatus were examined, they contained swimming trophonts whose characteristics revealed by Chatton–Lwoff stains describe a new species of Gymnodinioides. Its unique characteristic is the retention in the trophont of a polykinety similar to the falciform field of the tomite, the non-feeding, migratory microstome stage common to apostome genera. The trophont's anterior pattern of kineties also resembles that of the tomite. Speculations that speciation in Gymnodinioides may result from isolation on different hosts, changes in the internal feeding apparatus, delays or mishaps during morphogenesis, or habitat changes affecting the host must consider the case of G. pitelkae. The tomite may not respond to the signals from its premoult host to change to the macrostome infraciliature although the feeding apparatus is formed.     

Morphogenèse de Régénération chez le CiliéCondylostoma magnum (Spiegel): Etude ultrastructurale

SYNOPSIS Cortical events occurring in the course of regeneration in Condylostoma magnum (Spiegel) were studied by electron microscopy. The zone of regeneration is very rich in vacuoles and small vesicles formed from the plasma membrane. Multiplication of kinetosomes starts on the left side of kineties in the V-shaped left ventral area, normally implicated in stomatogenesis, at the level of the anterior kinetosomes of the somatic pairs. The proliferation proceeds by the appearance of young kinetosomes most often orthogonal to the old ones. This process of multiplication is very rapid and terminates in the formation of an “anarchic field” in which one observes that: (a) the newly formed kinetosomes do not possess all the associated postciliary fibers; and (b) when these fibers are detected, the kinetosomes are not in the same orientation. Differentiation of the adoral organelles takes place in the left part of the field (left primordium) by an alignment of the kinetosomes into 2 rows for each organelle (oriented perpendicularly to the antero-posterior axis of the ciliate), of which only one has the postciliary fibers. Ciliatogenesis occurs in numerous kinetosomes of the anarchic field; in certain kinetosomes it is achieved at the onset of their arrangement into organelles and is concomitant with growth of the nematodesmata. The 3rd (anterior) row of the organelles, the interkinetosomal desmata, and connections among neighboring organelles appear only secondarily. Differentiation of the paroral cilia occurs later. It takes place in the interior of the primordium, whose organization is primarily anarchic, and is accompanied by a progressive resorption of the major part of the newly formed kineties. Numerous kinetosomes of the right field have the associated postciliary fibers, which are not found at the level of the regenerated “polystichomonad” (paroral organization characteristic of C. magnum). Finally, the formation of new kinetosomes within a somatic kinety at the time of its elongation is described.     

Buccal and Somatic Infraciliature of Lagynus elegans (Engelmann, 1862) Quennersted, 1867 (Ciliophora, Prostomatea): Remarks on its Systematic Position

The somatic and buccal infraciliature of Lagynus elegans are described, and aspects of its division and conjugation are reported. Its somatic infraciliature is made up of 37–46 meridianal kineties composed of isolated kinetosomes that have thick and long kinetodesmal fibers. In the anterior zone of the cell, the circumoral infraciliature can be observed: it is composed of short, slightly oblique kinetal segments, which are formed of three kinetosomes each. The brosse of this species consists of 3 or 4 groups that possess 4 to 6 ciliated kinetosomes each; these kinetosomes lack kinetodesmal fibers. On the apical pole of the cell, surrounding the oral opening, a crown of nematodesmata is observed; these nematodesmata are connected to each other by a fibrillar structure. Taking into account these features, we propose that this genus be transferred from the order Prostomatida to a new family, Lagynidae, of the order Prorodontida.     

The Resorption of Cilia in Cyathodinium piriforme*

SYNOPSIS. The fine structure of the cilium, kinetosome, kinetodesmal fiber, and basal microtubules has been described in Cyathodinium piriforme. The ciliary axoneme is encased in an electron-dense jacket termed the axonemal jacket. This jacket surrounds the axoneme and is found midway between the axoneme and the ciliary membrane when viewed in cross section. Before division or reorganization the cilia are withdrawn into the cell. Intact cilia surrounded by their jackets are found in the cytoplasm during the early phases of retraction. Degradation of the axonemal microtubules precedes the dissolution of the axonemal jacket. Profiles of the jackets are observed after the microtubules have been resorbed. The cilia appear to detach from the kinetosomes. Barren kinetosomes are seen below the cell surface frequently with kinetodesmal fibers still attached. Whether all or some of these barren kinetosomes contribute to the formation of the new ciliary anlage cannot be ascertained.     

Cortical ultrastructure of Coleps bicuspis Noland, 1925 and the phylogeny of the class Prostomatea (Ciliophora)

The ultrastructure of Coleps bicuspis Noland, 1925 is described. The ciliate is a typical prostomate: the somatic kinetid is a monokinetid with a postciliary ribbon at triple 9, a kinetodesmal fibril originating near triplets 5, 6, 7 and an apparently radial transverse ribbon at triplet 4. The oral area is circular and has three brosse kineties associated with it. The brosse kineties are composed of dikinetids whose anterior kinetosome bears a tangential transverse ribbon and whose posterior kinetosome bears the fibrillar associates typical of a somatic monokinetid. The oral dikinetids are oriented parallel to the circumference of the oral cavity, which is surrounded by oral papillae and oral ridges. Pairs of nematodesmata, originating from oral dikinetid kinetosomes, are typically triangular in transection. A phylogeny of rhabdophoran ciliates is presented using the mixed parsimony algorithm and is discussed with reference to the systematic revisions of the phylum Ciliophora.