Ultrastructure of Prorodon (Ciliophora, Prostomatida) I. Somatic Cortex and Some Implications Concerning Kinetid Evolution In Prostomatid and Colpodid Ciliates

ABSTRACT. This study describes the ultrastructure of the somatic cortex of Prorodon aklitolophon and Prorodon teres. the meridionally arranged somatic kineties of both species can be separated into two parts: a short anterior part, which consists of a few somatic dikinetids (in which both kinetosomes are ciliated), and a longer posterior consisting of monokinetids. the somatic monokinetids are associated with a convergent postciliary microtubular ribbon, a transverse microtubular ribbon flatly inserted in front of the kinetosome, a short and steeply extending kinetodesmal fibre attached to kinetosomal triplet 5 and 7, and a desmose anterior to triplet 3. From this desmose, two to five prekinetosomal microtubules originate and extend anteriorly. the posterior kinetosome of the somatic dikinetids is associated with the same microfibrillar and microtubular structures as the somatic monokinetid, except that no prekinetosomal microtubules originate from the desmose. the anterior kinetosome has a single postciliary microtubule and a tangentially oriented transverse microtubular ribbon. the permanent collecting canals of the unique contractile vacuole system extend parallel and adjacent to the somatic kinetics of Prorodon . the collecting canals are supported by the prekinetosomal microtubules. A similarly organized contractile vacuole system is not yet known from any other ciliate group. One of the most surprising results of this investigation was finding a significant similarity between the somatic dikinetid pattern of Prorodon and the colpodid dikinetid pattern. A hypothesis is presented to illustrate the evolution of the somatic kinetid patterns in colpodid and prostomatid ciliates.     

The Ultrastructure of Chaenea teres and an Analysis of the Phylogeny of the Haptorid Ciliates

Chaenea teres has typical haptorid ultrastructure. The somatic monokinetid has two transverse microtubular ribbons, an overlapping postciliary microtubular ribbon, and a laterally directed kinetodesmal fiber. The evered cytopharynx forms a dome at the apical end of the cell. The base of the dome is surrounded by oral dikinetids. The left, anterior kinetosome of the oral pair is not ciliated and has a transverse microtubular ribbon, a nematodesmata and a single postciliary microtubule. The right, posterior kinetosome is ciliated and has only postciliary microtubules. The kinetosomes at the anterior ends of the somatic kinetics are close together and their transverse microtubules and nematodesmata contribute to the support of the cytopharynx. The transverse microtubules of these oralized somatic kinetosomes, together with those from the oral dikinetids, line the cytopharynx. Accessory or bulge microtubules arise perpendicular to the transverse microtubules. A dorsal brush of three kineties of clavate cilia is found on the cell surface just posterior to the oral region. Mucocysts and a single type of toxicyst are present. The toxicysts are confined to the oral region. There are multiple ovoid macronuclei that stain weakly. Micronuclei were not observed. Cladistic analysis indicates the Chaenea may be most closely related to Fuscheria and Acropisthium. The cladistic analysis also suggests that existing taxonomies of the subclass Haptoria need to be revised. We propose some modifications to Foissner & Foissner's classification that include transferring Helicoprorodon, Actinobolina, the buetschiliids, and the balantidiids to the order Haptorida and recognizing the close relationship between pleurostomes and spathidiids.     

A Reexamination of the Ultrastructure of Didinium nasutum and a Reanalysis of the Phylogeny of the Haptorid Ciliates

ABSTRACT. The cilia of Didinium nasutum are restricted to two girdles encircling the cell. Each row of cilia in both girdles is made up of two to three anterior pairs of kinetosomes followed by several single kinetosomes. Each single kinetosome has two sets of transverse microtubules, an overlapping postciliary microtubular ribbon, and a laterally directed kinetodesmal fiber. The pairs of kinetosomes are homologous to the oral dikinetids of other haptorians: the nonciliated kinetosome of the pair has a transverse microtubular ribbon that extends to line the membrane of the proboscis, a single short postciliary microtubule, and a nematodesma; the ciliated kinetosome has a ribbon of postciliary microtubules and two sets of transverse microtubules. The presence of these characters in Didinium invalidates Leipe & Hausmann's conclusion that the Didiniidae should be removed from the subclass that contains the other haptorians (Leipe, D. D. & Hausumann, K. 1989. Somatic infraciliature of the haptorid ciliate Homalozoon vermiculare (Kinetofragminophora, Gymnostomata) Ditransversalia n. subcl. and phylogenetic implications. J. Protozool. , 36 :280–289). In light of this, the justification for a subclass Ditransversalia is challenged and shown to be unnecessary.     

Light and Electron Microscopic Observations on the Heterotrich Ciliate Climacostomum virens

SYNOPSIS. Alveolar membranes and an epiplasm exist under the cell membrane of the noncontractile heterotrich ciliate Climacostomum virens. Postciliary microtubular ribbons join at the right of each somatic kinety to form a Km fiber. Two transverse microtubular fibers occur per kinetosomal pair. A myonemal network interconnects the kinetosomal bases intrakinetally and interkinetally. Ultrastructural comparisons are made between the contractile and noncontractile heterotrichs.
The buccal cortex consists of an adoral zone of membranelles, a peristomal field, a buccal tube, the apical membranelles, and a haplokinety. The kineties of the peristomal field and buccal tube are rows of paired kinetosomes, with a postciliary ribbon of microtubules arising from the posterior kinetosome of each pair, and a transverse ribbon and an oblique ribbon from the anterior kinetosome. No Km fibers exist in this region. The haplokinety is a collar of paired kinetosomes surrounding the cytostome; a postciliary microtubular ribbon descends from each kinetosomal pair into the cytostomal region. Ultrastructural details of the buccal cortex of C. virens and other heterotrichs are compared. The nemadesmata which lie under the membranelles are implicated in the body bending of C. virens.
Algae endosymbiotic in the cytoplasm of C. virens are described.     

Ultrastructural aspects of the somatic and buccal infraciliature ofColeps amphacanthus Ehrenberg 1833

Summary Somatic and buccal infraciliature ofColeps amphacanthus Ehrenberg 1833 were studied by light and electron microscopy. The somatic kineties are composed of monokinetids and 2 dikinetids at the anterior end of each kinety. The monokinetids are associated with postciliary microtubules at triplet 9, a kinetodesmal fiber at triplet 5 and 7 and nearly radially arranged transverse microtubules at triplet 4. The associated fibrillar systems of the posterior kinetosome of the dikinetids are like those of the monokinetids. The anterior kinetosome is associated with transverse microtubules at triplet 4 and one or few postciliary microtubules at triplet 9. The anterior kinetosome bears only a short cilium.The oral ciliature is composed of a kinety of nearly circumorally arranged paroral dikinetids and 3 adoral organelles at the ventral left side of the oral opening. Nematodesmata arising from the oral ciliature form the major component of the cytopharyngeal apparatus which is lined by microtubular ribbons of postciliary origin. The buccal cavity is surrounded by oral papillae which often contain toxicysts.     

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.     

Morphology and Ecology of Siroloxophyllum utriculariae (Penard, 1922) N. G., N. Comb. (Ciliophora, Pleurostomatida) and an Improved Classification of Pleurostomatid Ciliates

ABSTRACT. The morphology and infraciliature of Siroloxophyllum utriculariae (Penard, 1922) n. g., n. comb. were studied in live cells, with the scanning and transmission electron microscope, as well as in specimens impregnated with protargol and silver carbonate. The new genus, Siroloxophyllum , belongs to the Loxophyllidae and has a specific combination of characters, viz. an oral bulge surrounding almost the entire cell, three perioral kineties, a single brush kinety, and a single right dorsolateral kinety. The ecology and faunistics of S. utriculariae are reviewed. It is a rare and infrequent predator preferring clean freshwaters. The somatic monokinetid of S. utriculariae has typical haptorid ultrastructure, including two transverse microtubular ribbons. The oral bulge is patterned string-like with riffles containing the transverse microtubular ribbons originating from the oral kinetids. Perioral kineties 1 and 2 consist of dikinetids having one basal body each ciliated; the nonciliated basal body is associated with a nematodesmal and a transverse microtubular ribbon. Perioral kinety 3 consists of ciliated monokinetids having a fine structure similar to the somatic kinetids; they form triads with the dikinetids from perioral kinety 2. The classification of pleurostomatid ciliates is reviewed. Two suborders (Amphileptina, Litonotina) and three families (Amphileptidae, Litonotidae, Loxophyllidae n. fam.) are recognized and defined.     

Morphogenesis and Cortical Ultrastructure of Brooklynella hostilis,a Dysteriid Ciliate Ectoparasitic on Marine Fishes*

SYNOPSIS. Morphogenesis, and the cortical structures of Brooklynella hostilis, a cyrtophorine gymnostome ciliate ectoparasitic on marine fishes, were studied from protargol silver-impregnated preparations and with the aid of electron microscopy. The pattern of morphogenesis of Brooklynella is close to that found in less differentiated species of the families Chlamydodontidae (e.g., in the genus Trithigmostoma) and Dysteriidae (e.g., in the genus Hartmanula). The full number of kineties in the opisthe is restored after division from a segment of the left one of the 3 kinetics producing the oral rows. The oral rows consist of a double row of kinetosomes arranged in a zig-zag pattern; only the outer row is ciliated, the inner one being barren. However, the positions of the postciliary and transverse fibers indicate that the oral rows are not homologs of an undulating membrane but are akin to a membranelle. In association with the ventral somatic kinetosomes there are 4 postciliary fibers; a rather aberrant, transversally oriented kinetodesma; 2 microtubular, transverse fibers plus a transverse fibrousspur; and one to several ribbons of subkinetal microtubular fibers. Not directly associated with the kinetosomes are fibrous strands running subpellicularly between the kinetosomes and also deep into the cytoplasm. The cortical structures of Brooklynella are compared with those of some other groups of ciliates of about the same phylogenetic level in which the subkinetal microtubules can also be found– rhynchodine, suctorian, and chonotrich ciliates. The nasse consists of 6–8 nematodesmata not closely associated with the microtubular cytopharyngeal tube. The former have a distinctly developed densely fibrous capitulum containing barren kinetosomes which originally produced the nematodesma during stomatogenesis; the capitulum is connected by a fibrous link to the microtubular shaft. Extending from the oral rows to the capitula are fibrous structures strongly reminiscent of filamentous reticulum in hymenostome and peritrich ciliates. The structure of the posterio-ventral glandular organelle is also described and discussed.     

Ultrastructure of the Somatic and Buccal Cortex of the Tetrahymenine Hymenostome Glaucoma chattoni

SYNOPSIS The membranes, epiplasm, and fiber systems are described in the somatic cortex of Glaucoma chattoni strain HZ-1. Kinetodesmal fibers, postciliary and transverse microtubular ribbons, basal microtubules, transverse fibers and transverse accessory material are associated with kinetosomes. Longitudinal microtubular ribbons and mitochondria occur interkinetally. In the buccal cortex, the membranes, epiplasm and fibers of the 3 membranelles, the paroral kinety, the ribbed wall, and the cytostome are described. Comparisons between G. chattoni and other ciliates reveal ultrastructural differences of possible systematic significance. In the somatic cortex of this and other tetrahymenines. Iongitudinal microtubular ribbons and basal microtubules occur concurrently. In the buccal cortex, alveoli are absent in tetrahymenine membranelles. A table is presented of the fiber systems associated with single somatic kinetosomes of various ciliates whose cortical ultrastructure has been studied to date.     

Description of the Infraciliature and Morphogenesis in the Ciliate Urotricha ondina N. Sp. (Prorodontida, Urotrichidae)

Recent works on prostomatid ciliates show that some genera of this group have a differentiated oral infraciliature and that their stomatogenesis during division involves the proliferation of only a few somatic kineties. These findings have significant implications regarding the iaxonomic status of these genera and also on the terminology used for the oral structures. In Urotricha ondina , the oral infraciliature consists of (1) a paroral kinety formed of paired kinetosomes that encircle the cytostome at the anterior pole of the cell and (2) 3 adoral organelles, each formed of 2 rows of kinetosomes, ventral in position and obliquely disposed, lying above 3 short somatic kineties that do not reach the anterior pole of the cell. This oral ciliature —formerly known as the corona and brosse, respectively—originate during stomatogenesis from the proliferation of 4 somatic kineties that lie posterior to the adoral organelles of the parental cell.     

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.     

Fine Structure and Molecular Phylogeny of Parametopidium circumlabens (Ciliophora: Armophorea), Endocommensal of Sea Urchins

Metopid armophoreans are ciliates commonly found in anaerobic environments worldwide; however, very little is known of their fine structure. In this study, the metopid Parametopidium circumlabens (Biggar and Wenrich 1932) Aescht, 1980, a common endocommensal of sea urchins, is investigated for the first time with emphasis on transmission electron microscopy, revealing several previously unknown elements of its morphology. Somatic dikinetids of P. circumlabens have a typical ribbon of transverse microtubules, an isolated microtubule near triplets 4 and 5 of the anterior kinetosome, plus two other microtubules between anterior and posterior kinetosomes, a short kinetodesmal striated fiber and long postciliary microtubules. In the dikinetids of the perizonal stripe, the kinetodesmal fiber is very pronounced, and there is a conspicuous microfibrillar network system associated with the kinetosomes. A new structure, shaped as a dense, roughly cylindrical mass surrounded by microtubules, is found associated with the posterior kinetosome of perizonal dikinetids. The paroral membrane is diplostichomonad and the adoral membranelles are of the “paramembranelle” type. Bayesian inference and maximum‐likelihood analysis of the 18S‐rDNA gene unambiguously placed P. circumlabens as sister group of the cluster formed by ((Atopospira galeata, Atopospira violacea) Metopus laminarius) + Clevelandellida, corroborating its classification within the Metopida.     

Etude Ultrastructurale du CiliéKuklikophrya dragescoi gen. n., sp. n.*

SYNOPSIS The cortical infraciliature of Kuklikophrya dragescoi gen. n., sp. n. is composed of double kinetosomes. Each kinetosome has transverse fibers. The anterior transverse fibers are associated with a sheet of dense material and the posterior transverse fibers are directed toward the posterior part of the body. The posterior kinetosome of a pair has only a short protuberance in the position of the kinetosomal fiber. The cortex has a well developed alveolar layer and a thick ecto-endoplasmic boundary. A distinctive characteristic of the buccal ciliature is the circumoral ciliature whose infraciliature is made up of pairs of cilia-bearing kinetosomes. The antero-posterior polarity of the paroral segment is in inverse relationship to that of the remaining ciliature of the organism. The adoral and preoral ciliary organelles consist of 2 rows of kinetosomes, each of which bears postciliary fibers. A frame of nematodesmata surrounds the cytopharynx which is supported by microtubular bands which impart to it a very specific laminated appearance. The “phagoplasm” is formed by “vermicelli”-like vesicles. The micronucleus is found in the perinuclear area of the macronucleus.     

Somatic Infraciliature in the Haptorid Ciliate Homalozoon vermiculare (Kinetophragminophora, Gymnostomata) Ditransversalia N. Subcl. and Phylogenetic Implications

The ultrastructure of the cortex of Homalozoon vermiculare is described. The ventral side bears 13–15 iongitudinal kineties composed of monokinetids. On the dorsal surface, there are 3 kineties, 2 of which are composed of dikinetids in the anteriormost part of the cell. Consequently there exist 3 different kinds of kinetids within the somatic cortex: 1) The monokinetids on the ventral side are associated with a kinctodesmal fibril, 2 transverse microtubular ribbons and 7 postciliary microtubules in a double-row configuration; 2) The monokinetids on the dorsal side are very similar but they are associated with just 3 very 'short postciliary microtubules; 3) The posterior kinetosome of the dorsal dikinetids bears the same fibrillar associates as the dorsal monokinetid, but it lacks the second transverse ribbon. The anterior kinetosome of each pair is associated with a single postciliary' microtubule. The kinetid organization of Homalozoon is compared to that of other members of the Haptorida. Their phylogeny is discussed. A monophyleiic taxon within the litostomate ciliates is characterized by data on the somatic kinetids, and the new subclass Ditransversalia n. subcl. is constituted. The new subclass comprises the genera Balantidium, Bryophyllum, Enchelydium, Homalozoon, Isotricha, Lacrymaria, Lepidolrachelophyllum, Spathidium and Vestibulongum .     

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 Ultrastructure of Zosterodasys agamalievi (Ciliophora: Synhymeniida)

ABSTRACT. The cell surface of the synhymeniid ciliate, Zosterodasys agamalievi , consists of shallow kinetal grooves separated by low cortical ridges. Numerous electron-opaque bodies are located in the cortical ridges, inside the kinetal grooves, and are distributed in parallel rows between adjacent kineties. Well-developed alveoli are present beneath the cell surface membrane. Zosterodasys agamalievi has a single micronucleus and a homomerous macronucleus. The infraciliature of the somatic monokinetid consists of an anteriorly-directed kinetodesmal fiber, a well-developed divergent postciliary microtubular ribbon, radially-oriented transverse microtubules, and a short striated rootlet, which extends anteriorly from the base of the kinetosome into the cell. Zosterodasys agamalievi has a perioral band of paired cilia, the synhymenium, that winds obliquely across the ventral surface of the body, just posterior to the cytostome. The infraciliature of the anterior kinetosome of the synhymenium consists of two postciliary microtubules; a well-developed, divergent post-ciliary ribbon of microtubules and a short kinetodesmal fiber are associated with the posterior kinetosome. The cytopharynx is supported by 14-16 nematodesmata which are capped distally by a capitulum. The cytopharynx is bound proximally by a fibrous sheath and is lined by radially-arranged microtubular ribbons. No obvious oral ciliature is present.     

Oral Monokinetids in the Free-Living Haptorid Ciliate Enchelydium polynucleatum (Ciliophora,Enchelyidae): Ultrastructural Evidence and Phylogenetic Implications1

Special ultrastructural characteristics of the haptorid soil ciliate Enchelydium polynucleatum Foissner, 1984 are the restriction of the parasomal sacs to the area of the “brush” and finger-like projections of the food vacuole membrane into the lumen of the vacuole. The general organization of the infraciliature is similar to that of Spathidium and some buetschliids because the anterior ends of the somatic kineties are condensed and obliquely bent. Enchelydium is similar to haptorids and buetschliids in possessing monokinetid somatic fibrillar structures with the classical fibrillar associates: 1) a short kinetodesmal fiber; 2) two transverse microtubular ribbons; 3) a long postciliary microtubular ribbon; and 4) a system of overlapping subkinetal microtubules, which seems to be absent in the buetschliids. Unlike Spathidium and all other haptorids so far investigated ultrastructurally, serial sections show that there are no oral dikinetids, as in the endocommensal buetschliids and balantidiids. Instead, three to six anterior kinetids in each ciliary row have nematodesmal bundles extending into the cytoplasm and surrounding the cytopharynx. These kinetids lack cilia and all fibrillar associates except enlarged transverse ribbons, which extend anteriorly and inwards to support the cytopharynx. Other similarities between the buetschliids and Enchelydium are the conspicuous rough endoplasmic reticulum and abundant sausage-like vesicles in the oral region. As in other haptorids, Enchelydium has two types of toxicysts and one type of mucocyst. These observations strongly suggest that Enchelydium belongs to the ancestral stock of both the Haptorida and the Archistomatida. The similarities in the somatic and oral infraciliature and ultrastructure of the Haptorida and the Archistomatida suggest that they belong to the same subclass, Haptoria Corliss, 1974.     

Light and Electron Microscopical Observations on the Stomatogenesis of the Ciliate Coleps amphacanthus Ehrenberg, 1833

Light and electron microscopical observations on the stomatogenesis of Coleps amphacanthus Ehrenberg, 1833, show that this "gymno"-stome ciliate has a well developed oral ciliature made of 19–23 "paroral dikinetids" and three "adoral organelles." These structures were previously known as "circumoral ciliature" and "dorsal brosse," and it was thought that they originated from the distal ends of all the 22–26 somatic kineties. Contrary to this view, only four stomatogenic kineties (K1, Kn, Kn-1, and Kn-2) are involved in stomatogenesis of the opisthe. All paroral dikinetids arise from one single kinetofragment (KF1) to the right of the oral anlage while the adoral organelles originate from the three left kinetofragments (KFn, KFn-1, and KFn-2). In particular, the future paroral dikinetids perform a complex morphogenetic movement that leads to a situation where the postciliary microtubules of the once posterior kinetosome of each oral dikinetid give rise to the cytopharyngeal microtubular ribbons. The postciliary origin of the cytopharyngeal ribbons which could only be detected by an EM study of stomatogenesis shows that the basket of Coleps belongs to the cyrtos-type and not to the rhabdos-type basket, where transverse microtubules accompany the basket-forming nematodesmata. The taxonomic implications of these observations, which may lead to a revision of the systematic position of the genus Coleps , are discussed.     

Cortical ultrastructure of the Scuticociliates Dexiotricha media and Dexiotricha colpidiopsis (Hymenostomata).

Cortical ultrastructure of the scuticociliates Dexiotricha media and Dexiotricha colpidiopsis was investigated. The following elements of the somatic cortex were studied: the cell membrane, alveolar membranes and the epiplasm, kinetodesmal fibers, postciliary and transverse microtubular ribbons, and transverse fibers associated with single and paired kinetosomes; mitochondria and single microtubules located in interkinetal ridges; mature and early extrusion stages of mucocysts; the expulsion vacuole pore and tube, the nephridioplasm and the cytoproct. In the buccal cortex, the paroral kinety-ribbed wall complex, the 3 polykineties, and the cytostome-cytopharynx were investigated. Comparative survey of ciliate ultrastructure indicates 2 principal orientation patterns for kinetodesmal and postciliary fibers, recognition of which leads to reevaluation of the theory of paroral kinety formation and the ideas of homology based on this theory. Ultrastructurally, the scuticociliates are not distinct from tetrahymenines and peniculines; the 3 groups appear to be 1 assemblage.     

THE ORGANIZATION AND EVOLUTION OF MICROTUBULAR ORGANELLES IN CILIATED PROTOZOA

(1) Ciliated protozoa are viewed as unicellular organisms structured in a hierarchy of organizational levels that include the macromolecular, suborganellar, unit organellar, organellar complex, and organellar system. (2) The ciliate cortex is divided into two major functional regions, the somatic region and the oral region. The fundamental component of the cortex is an organellar complex, the kinetid, whose organizing centre is the kinetosome with which are associated three fibrillar associates diagnostic of ciliates. These three fibrillar associates are the periodically striated kinetodesmal fibril and two microtubular ribbons, the transverse and postciliary ribbons. (3) Somatic and oral kinetids are found to be of three major types: monokinetids are composed of one kinetosome and its fibrillar associates; dikinetids are composed of two kinetosomes and their fibrillar associates; polykinetids are composed of more than two kinetosomes and their fibrillar associptes. (4) The mechanisms underlying kinetid function and development remain largely unexplored. Research into the molecular biology and ultrastructure, especially of mutant forms, should provide basic insights in the near future. (5) The conservation of kinetid structure across major phyla of organisms suggests that this subcellular structure should be useful in phylogenetic analysis despite the concepts of ‘chemical identity’ and ‘organic design’. (6) The evolutionary rate of change of oral features is greater than that of somatic features, probably due to developmental and ecological factors. Nevertheless both cortical regions are constrained by the phenomenon of structural conservatism; that is, the conservation of structure through time is inversely related to the level of biological organization. (7) Eight major groupings of ciliate species are recognized, based on ultrastruc-tural features of the cortex. Several examples of differences between these eight groups and the groups presently recognized are discussed.