Evolution and Diversity of Amitochondrial Zooflagellates

ABSTRACT. The amitochondrial flagellates comprise the Archamoebae, the Metamonada (orders: Retortamonadida, Diplomonadida, Oxymonadida) and the Parabasala. Molecular rDNA sequence comparison has shown that the diplomonads are very ancient and placed at the base of the tree, but the position of the parabasalids before or after the Euglenozoa and other mitochondriate protists is not definitively determined and such molecular data are required to place the other groups. Common cytological features such as one basal body and a basal body-nucleus connector show that the mastigamoebids or Archamoebae are an evolutionary lineage. The metamonad grouping is heterogenous; the three orders have in common two pairs of basal bodies, no Golgi and could be poly- or paraphyletic. The Parabasala is an evolutionary lineage with a large set of homologous cytological characters. A biochemical study of the cytoskeletal structures such as costa, parabasal fibre, preaxostylar fibre and undulating membrane in trichomonads reveals a large diversity in the protein composition among the genera that fits with the large distance between the taxa estimated by rDNA sequencing. Molecular comparison in seven tnchomonad genera indicates that the Trichomonadinae and Tritrichomonadinae form two divergent branches, the genera Hypotrichomonas and Monocercomonas are now incorporated in the tritrichomonad branch, and Trichomitus is at the bifurcation of the two branches. This scheme is different from the classic tree but not incompatible with it.     

Lightmicroscopic Observations on Structure and Division of Histomonas meleagridis (Smith)*

SYNOPSIS. Culture forms and lumen-dwelling phases of the ameboflagellate Histomonas meleagridis, which are structurally indistinguishable from each other, have a single flagellum. Their well-developed pelta is connected to the anterior segment of the broad, spatulate axostylar capitulum, applied to the left-ventral surface of the nearly spheroid or somewhat ellipsoid or ovoid nucleus. The capitulum narrows into a very slender axostylar trunk that tapers to a fine point and does not project beyond the body surface. The parabasal apparatus consists of a V-shaped parabasal body and a large parabasal filament. A new flagellum appears early during division and soon approaches its full length. The 2 flagella persist thruout division and each becomes the locomotory organelle of a daughter histomonad. The arms of the parental parabasal body appear to separate, each going to 1 of the daughter mastigont systems; some parabasal material is lost early in division. The 2nd arm is regenerated in each daughter parabasal body. The large parabasal filament seems not to be retained in the parental mastigont system, and new filaments are seen at both poles before 2 daughter nuclei are formed. The old axostyle degenerates from the anterior toward the posterior end; at the same time lamellar primordia of the daughter pelta-axostyle complexes appear in the separating mastigont systems that are connected by an extranuclear spindle during the entire division process. The structure and taxonomic status of H. meleagridis are discussed in the light of this and previous studies.     

The flagellar apparatus and cytoskeleton of the dinoflagellates

Summary Modern microscopical approaches have allowed more accurate investigations of the three-dimensional nature of the dinoflagellate flagellar apparatus (FA) and several other cytoskeletal protein complexes. Our presentation overviews the nature of the dinoflagellate FA and cytoskeleton in a number of taxa and compares them with those of other protists. As with other protists, the FA of the dinoflagellates can be characterized by the presence of fibrous and microtubular components. Our studies and others indicate that the dinoflagellate FA can be expected to possess a striated fibrous root on the basal body of the transverse flagellum and a multimembered microtubular root on the basal body of the longitudinal flagellum. Two other features that appear widespread in the group are the transverse striated root associated microtubule (tsrm) and the transverse microtubular root (tmr). The tsrm extends at least half the length of the transverse striated root while the tmr extends from the transverse basal body toward the exit aperture of the transverse flagellum. In most cases, the tmr gives rise to several cytoplasmic microtubules at a right angle. The apparent conserved nature of these roots leads us to the conclusion that the dinoflagellate FA can be compared to the FA of the cryptomonads, chrysophytes, and the ciliates for phylogenetic purposes. Of these groups, the chrysophytes possess an FA with the most structures in common with the dinoflagellates. Our immunomicroscopical investigations of the microtubular, actin and centrin components of the dinoflagellate cytoskeleton point to the comparative usefulness of these cytological features.Abbreviations aptb apical transverse microtubular band - FA flagellar apparatus - Imr longitudinal microtubular root - mls multilayered structure - tmr transverse microtubular root - tmre transverse microtubular root extension - tsr transverse striated fibrous root - tsrm transverse striated root associated microtubule     

Ultrastructure of the genus Enteromonas da Fonseca (Zoomastigophorea) and revision of the order of Diplomonadida Wenyon]

Fine structure of 2 species of Enteromonas, one from the intestine of the salamander, Triturus vulgairs, and another from the feces of domestic rabbit, Oryctolagus cuniculi, is described. The pyriform cell has an anteriorly located nucleus. The 4 flagella originate from an area near the anterior end of the nucleus. The recurrent flagellum (R) is lodged in a ventral depression or cytostome. The kinetosomes, arranged into 2 pairs, anterior (no. 1, no. 2) and posterior ("3, R), are interconnected by microfibrils. One microtubular fiber, connected to kinetosome "1, is situated near the anterior surface of the nucleus. Another, subnuclear, microtubular fiber is homologous to the "crossed" fiber found in Diplozoa. The cytostome is bordered by 2 lips: the preeminent left lip is equipped with several rows of microtubules, while the right lip contains only a thin microtubular fiber associated with microfibrils. The cytostome occupies 2/3 of the ventral surface. The recurrent flagellum passos over the anterior surface of the cell and then comes to lie in the cytostome. The bacteria are phagocytosed in the bottom part of the cytostome between the 2 distended lips. They are digested in numerous vacuoles. The undigested residual bodies are evacuated by a rupture of the cell membrane. The ergastoplasm is concentrated near the cell periphery. Mitochondria and the Golgi apparatus are absent. In the cyst stage, the multinucleate cell is enclosed in a microfibrillar membrane; the axonemes lie free in the cytoplasm. Diplomonad forms of Enteromonas resembling Hexamita are numerous, except that the cytostome is different in these 2 genera. In such forms, the arrangement of the 2 individuals often has binary axial symmetry, but on occasion they are associated in a more anarchic fashion. The mastigont of Enteromonas is organized like that of a single zooid of a diplozoon. It is possible that the genus Enteromonas is ancestral to Diplomonadida and that the diplomonad state, transitory in Enteromonas, became permanently established in Diplomonadida. Enteromonas appears to be more primitive than the other genera of Diplomonadida. Thus we propose 2 suborders: Enteromonadina, subord. nov. with the genus Enteromonas, and Diplomonadina Wenyon, emend., with the genera Trepomonas, Trigonomonas, Hexamita, Spironucleus, Octomitus, Giardia. The arrangement of the kinetosomes and the existence of a cytostome are the principal characters common to Enteromonas and Retortamonadida, while their "accessory" fibers are not homologous. A more complete study of division of the 2 zooflagellate orders is necessary for the presentation of a more detailed evolutionary scheme of these groups.     

Ultrastructure of Ditrichomonas honigbergii N. G., N. Sp. (Parabasalia) and Its Relationship to Amitochondrial Protists

ABSTRACT. Ditrichomonas honigbergii n. g., n. sp. is a small trichomonad flagellate that has three emergent flagella arising from four basal bodies, a parabasal apparatus (single dictyosome with associated striated flagellar rootlets), a microtubular axostyle, a short undulating membrane, and hydrogenosomes. Cultures of D. honigbergii were isolated from the sediments of a freshwater lake and there is no known metazoan host. Cells form walled cysts with internalized flagella and go through all phases of the life cycle (excystment, binary division, encystment) without any perturbations to the culture medium. Ditrichornonas honigbergii is capable of ingesting and digesting bacteria by phagocytosis. These facts suggest that D. honigbergii may be a free-living inhabitant of oxygen-reduced environments. The structure of D. honigbergii is similar to that of retortamonads and the relationship of trichomonads to other amitochondrial protists is discussed.     

Transformation of the flagella and associated flagellar components during cell division in the coccolithophoridPleurochrysis carterae

Summary Immunofluorescence microscopy, conventional and high voltage transmission electron microscopy were used to describe changes in the flagellar apparatus during cell division in the motile, coccolithbearing cells ofPleurochrysis carterae (Braarud and Fagerlund) Christensen. New basal bodies appear alongside the parental basal bodies before mitosis and at prophase the large microtubular (crystalline) roots disassemble as their component microtubules migrate to the future spindle poles. By prometaphase the crystalline roots have disappeared; the flagellar axonemes shorten and the two pairs of basal bodies (each consisting of one parental and one daughter basal body) separate so that each pair is distal to a spindle pole. By late prometaphase the pairs of basal bodies bear diminutive flagellar roots for the future daughter cells. The long flagellum of each daughter cell is derived from the parental basal bodies; thus, the basal body that produces a short flagellum in the parent produces a long flagellum in the daughter cell. We conclude that each basal body in these cells is inherently identical but that a first generation basal body generates a short flagellum and in succeeding generations it produces a long flagellum. At metaphase a fibrous band connecting the basal bodies appears and the roots and basal bodies reorient to their interphase configuration. By telophase the crystalline roots have begun to reform and the rootlet microtubules have assumed their interphase appearance by early cytokinesis.Abbreviations CR1, CR2 crystalline roots 1 and 2 - CT cytoplasmic tongue microtubules - DIC differential interference contrast light microscopy - H haptonema - HVEM high voltage transmission electron microscopy - IMF immunofluorescence microscopy - L left flagellum/basal body - M metaphase plate - MT microtubule - N nucleus - R right flagellum/basal body - R1, R2, R3 roots 1, 2, and 3 - TEM transmission electron microscopy     

The cytology and ultrastructure of zoospores of Hydrurus foetidus (Chrysophyceae)

The unusual tetrahedral shape of Hydrurus foetidus (Vill.) Trev. zoospores is associated with a complex skeletal system of microtubules extending from a broad flagellar root (up to 19 microtubules) into each of three, pointed anterior processes. The posterior end, also pointed and supported by a separate set of microtubules, contains a single large chloroplast with a prominent posterior furrow containing mitochondrial elements. A large immersed pyrenoid is penetrated by paired thylakoids. There is no eyespot. Numerous large Golgi bodies occur immediately anterior to the nucleus and up to 5–6 contractile vacuoles lie near the cell surface at the anterior end. Two terminally inserted flagella extend from the cell surface, a long one serving for cell locomotion, and the other vestigial with an axonemal pattern of 9+0. The flagellar root system consists of: (1) a thin, striated rhizoplast extending from the basal body of the long flagellum and ramifying over the surface of a conspicuous, anteriorly directed, conical projection of the nucleus; (2) a broad microtubular root which emanates from near the basal body of the long flagellum and appears to function as a MTOC; (3) a compound root, consisting of a striated fiber and two associated microtubules, which runs alongside the basal body of the stubby flagellum before terminating at the cell surface; and (4) a short two-membered microtubular root, also associated with the basal body of the stubby flagellum. Other components of the flagellar apparatus include a large dense body near the proximal end of the basal body of the short flagellum, and a small, dense, core-like structure closely associated with one of its triplet fibers. The flagellar apparatus of H. foetidus is remarkably similar in ultrastructure to that of Chrysonebula holmesii Lund.     

Malawimonas jakobiformis n. gen., n. sp. (Malawimonadidae n. fam.): A Jakoba-like Heterotrophic Nanoflagellate with Discoidal Mitochondrial Cristae

Malawimonas jakobiformis n. gen., n. sp., is established for a bacterivorous heterotrophic nanoflagellate isolated from the Malawi shore of Lake Nyasa (eastern Africa). Trophic stages observed were anteriorly biflagellate and naked. The posterior flagellum of a trophic cell resided in a conspicuous groove on the ventral surface, and bore a prominent vane. A Golgi stack and a mitochondrion with discoidal cristae were present anterior to the nucleus. The kinetid consisted of two short, slightly separated basal bodies, four microtubular roots, and associated fibers and bands. The three microtubular roots associated with the posterior basal body were associated with the ventral groove, while the single root associated with the anterior basal body gave rise to secondary cytoskeletal microtubules. Dividing cells became rounded, with persistent flagella. Cysts were uninucleate, and had thin organic walls without clearly differentiated apertures or ornamentation but with conspicuous attachment pads. Kinetid elements were present within cysts. On the basis of microscopical features, especially those of the kinetid, the nearest relatives of M. jakobiformis are the mitochondriate “jakobid” protists (families Histionidae and Jakobidae) and the amitochondriate retortamonads. Malawimonadidae n. fam. is established to accommodate this species.     

The ultrastructure of Ancyromonas, a eukaryote without supergroup affinities

The small heterotrophic flagellate Ancyromonas (=Planomonas) lacks close relatives in most molecular phylogenies, and it is suspected that it does not belong to any of the recognized eukaryote 'supergroups', making it an organism of great evolutionary interest. Proposed relatives include apusomonads and excavates, but limited understanding of the ancyromonad cytoskeleton has precluded identification of candidate structural homologies. We present a detailed analysis of the ultrastructure of Ancyromonas through computer-based reconstruction of serial sections. We confirm or extend previous observations of its major organelles (mitochondria, Golgi body, extrusomes, etc.) and pellicle, and distinguish a system of stacked endomembranes that may be developmentally connected to the glycocalyx. Ancyromonas has two basal bodies, each with its own flagellar pocket. The anterior basal body associates with two microtubular elements: a doublet root that runs from between the basal bodies to support the cell's rostrum, and a short singlet root. The posterior basal body is associated with two multi-microtubular structures and a singlet root. One multi-microtubular structure, L1, is a conventional microtubular root. The other structure appears as a curved ribbon of ～8 microtubules near the basal body, but then flares out into two multi-microtubular elements, L2 and L3, plus two single microtubules. The posterior singlet root originates independently near this second complex. L1, the singlet, L2, and L3 all support the posterior flagellar pocket and channel. We also identified several groups of peripheral microtubules. Possible homologies with the flagellar apparatus of both apusomonads and excavates include a splitting root on the right side of the posterior basal body and a singlet root, both supporting a longitudinal channel or groove associated with the posterior flagellum. The anterior flagellar apparatus in each includes a root supporting structures to the left of the anterior flagellum. Given the probable deep divergences of Ancyromonas, apusomonads and excavates within eukaryotes, it is possible that the eukaryotic cenancestor also possessed these features.     

The flagellar root system inHeterosigma akashiwo (Raphidophyceae)

Summary The flagellar apparatus of 3 isolates ofHeterosigma akashiwo (Hada) Hada has been studied by serial sectioning. The two basal bodies lie at almost right angles to one another, but in a different plane, and are interconnected by an extensive root system. This consists of three roots (i) a massive cross-banded fibrous root (= rhizoplast) which extends from near the proximal ends of both basal bodies to the anterior surface of the nucleus, (ii) a compound microtubular root with a layered structure, associated with the hairy anterior flagellum and extending to the anterior surface and (iii) the rhizostyle which passes between the two basal bodies leading anteriorly to a vesicle in the flagellar groove region and following the nucleus posteriorly terminating deep in the cytoplasm. Both the characteristic arrangement of the basal bodies and the presence of the complex layered structure are characteristic of theRaphidophyceae. The broad microtubular root, however, to which the layered structure is attached, appears to be characteristic of nearly all heterokont algae, fungi and protozoa so far examined. Thus, our findings have important implications on phylogenetic relationships within the heterokonts and lead us to question whether some of the present classes such as theChrysophyceae andXanthophyceae are indeed natural groups.     

Ultrastructure of the flagellar apparatus inPleurochrysis (classPrymnesiophyceae)

Summary The ultrastructure of the flagellar apparatus of aPleurochrysis, a coccolithophorid was studied in detail. Three major fibrous connecting bands and several accessory fibrous bands link the basal bodies, haptonema and microtubular flagellar roots. The asymmetrical flagellar root system is composed of three different microtubular roots (referred to here as roots 1,2, and 3) and a fibrous root. Root 1, associated with one of the basal bodies, is of the compound type, constructed of two sets of microtubules,viz. a broad sheet consisting of up to twenty closely aligned microtubules, and a secondary bundle made up of 100–200 microtubules which arises at right angles to the former. A thin electron-dense plate occurs on the surface of the microtubular sheet opposite the secondary bundle. The fibrous root arises from the same basal body and passes along the plasmalemma together with the microtubular sheet of root 1. Root 2 is also of the compound type and arises from one of the major connecting bands (called a distal band) as a four-stranded microtubular root and extends in the opposite direction to the haptonema. From this stranded root a secondary bundle of microtubules arises at approximately right angle. Root 3 is a more simple type, composed of at least six microtubules which are associated with the basal body. The flagellar transition region was found to be unusual for the classPrymnesiophyceae. The phylogenetic significance of the flagellar apparatus in thePrymnesiophyceae is discussed.     

Ultrastructure of Trimastix pyriformis (Klebs) Bernard et al.: similarities of Trimastix species with retortamonad and jakobid flagellates.

Trimastix pyriformis (Klebs 1893) Bernard et al. 1999, is a quadriflagellate, free-living, bacterivorous heterotrophic nanoflagellate from anoxic freshwaters that lacks mitochondria. Monoprotist cultures of this species contained naked trophic cells with anterior flagellar insertion and a conspicuous ventral groove. Bacteria were ingested at the posterior end of the ventral groove, but there was no persistent cytopharyngeal complex. The posterior flagellum resided in this groove, and bore two prominent vanes. A Golgi body (dictyosome) was present adjacent to the flagellar insertion. The kinetid consisted of four basal bodies, four microtubular roots, and associated fibers and bands. Duplicated kinetids, each with four basal bodies and microtubular root templates, appeared at the poles of the open mitotic spindle. Trimastix pyriformis is distinguishable from other Trimastix species on the basis of external morphology, kinetid architecture and the distribution of endomembranes. Trimastix species are most similar to jakobid flagellates, especially Malawimonas jakobiformis, and to species of the retortamonad genus Chilomastix. Retortamonads may have evolved from a Trimastix-like ancestor through loss of "canonical" (easily seen with electron microscopy) endomembrane systems and elaboration of cytoskeletal elements associated with the cytostome/cytopharynx complex.     

ULTRASTRUCTURAL STUDIES ON BIGELOWIELLA NATANS, GEN. ET SP. NOV., A CHLORARACHNIOPHYTE FLAGELLATE

Three isolates from the Provasoli-Guillard National Center for Culture of Marine Phytoplankton at Bigelow Laboratory, previously labeled Pedinomonas sp. and Pedinomonas minutissima from the green algal class Pedinophyceae, have been examined by light microscopy and TEM and shown to belong to the Chlorarachniophyceae, a class of nucleomorph-containing amebae. The three isolates represent the first chlorarachniophycean flagellates to be discovered. The ultrastructure of the cells has been examined in detail, with particular emphasis on the flagellar apparatus, a feature not examined in detail in chlorarachniophytes before. Cells are basically biflagellate, but the second flagellum is represented by a very short basal body only. Flagellar replication has shown this flagellum to be the mature stage, that is, the no. 1 flagellum, whereas the long emergent flagellum is the no. 2 flagellum that shortens into a short basal body during cell division. Mitosis is open with a pair of centrioles at each pole. Emergent flagella are absent during mitosis. Cells may form cysts, and the flagellar basal bodies and part of the flagellar roots are maintained in the cysts. Four microtubular roots emanate from the basal bodies, and the path of one of them is very unusual and very unlike any other known flagellate. No striated roots were observed. Other fine-structural features of the cell include a very unusual type of pyrenoid and a special type of extrusome. Cells are mixotrophic. The three isolates are very similar and are described as Bigelowiella natans , gen. et sp. nov. Ultrastructurally, chlorarachniophytes do not show close relationship to any known group of algae or other protists.     

DNA fluorescent stain accumulates in the Golgi but not in the kinetosomes of amitochondriate protists.

Hindgut symbiotic trichomonads (uninucleate Caduceia versatilis, and multinucleate Stephanonympha sp. and Snyderella tabogae) from the dry-wood-eating termite Cryptotermes cavifrons (Kalotermitidae) accumulate DAPI (4,6diamidino-2-phenylindole) in the membranous sacs of the Golgi complex. This form of Golgi complex, typical of protists in the class Parabasalia, is called a parabasal body. Trichomonads contain organellar systems, mastigonts, that consist of four undulipodia (e.g. eukaryotic flagella and cilia), axostylar microtubules, a parabasal body and other structures. These cells bear from one (in the case of Caduceia) to hundreds (in the case of Snyderella) of mastigonts. These features are characteristic of their protist class (Parabasalia). The nuclei of all three species stained with DNA-specific stains: DAPI, SYTOX, acridine orange, propidium iodide, ethidium bromide and Feulgen, at optimal concentrations, but kinetosomes failed to stain at all. The nuclei, parabasal bodies and symbiotic bacteria (but no microtubular structures) fluoresced in glutaraldehyde-fixed cells stained with 1.45 microM DAPI. Parabasal bodies of Snyderella and Caduceia treated to remove lipids with Triton X-100, or treated with 5% trichloroacetic acid, lacked DAPI-fluorescence. I conclude that DNA, present as expected in nuclei and bacterial symbionts, is absent from and not associated with calonymphid kinetosomes. The reason for DNA-RNA stain accumulation in the Golgi cistemae is not clear.     

Vischeria stellata (Eustigmatophyceae): ultrastructure of the zoospores,with special reference to the flagellar apparatus

Summary Ultrastructure of the zoospores ofVischeria stellata (R. Chodat ex Poulton) Pascher is investigated, with particular reference to the system of flagellar roots. Microtubular roots and a rhizoplast are present and a model showing their distribution is proposed. Four microtubular roots attach to the basal bodies in a system basically similar to that displayed by the heterokont algae and fungi. The rhizoplast is also similar to that of other heterokont algae. We conclude from these observations that the class Eustigmatophyceae should be placed within the division Heterokontophyta.Abbreviations C chloroplast - B basal body of the emergent flagellum - B' second basal body - E eyespot - F emergent flagellum - FS flagellar swelling - LV lamellate vesicle - M mastigonemes - MTs microtubules - N nucleus - R 1–R 4 microtubular roots - Rh rhizoplast - SB striated band - SV spiral vesicle     

Cell division of Giardia intestinalis: flagellar developmental cycle involves transformation and exchange of flagella between mastigonts of a diplomonad cell

Giardia intestinalis is a binucleated diplomonad possessing four pairs of flagella of distinct location and function. Its pathogenic potential depends on the integrity of a complex microtubular cytoskeleton that undergoes a profound but poorly understood reorganization during cell division. We examined the cell division of G. intestinalis with the aid of light and electron microscopy and immunofluorescence methods and present here new observations on the reorganization of the flagellar apparatus in the dividing Giardia. Our results demonstrated the presence of a flagellar maturation process during which the flagella migrate, assume different position, and transform to different flagellar types in progeny until their maturation is completed. For each newly assembled flagellum it takes three cell cycles to become mature. The mature flagellum of Giardia is the caudal one that possesses a privileged basal body at which the microtubules of the adhesive disk nucleate. In contrast to generally accepted assumption that each of the two diplomonad mastigonts develops separately, we found that they are developmentally linked, exchanging their cytoskeletal components at the early phase of mitosis. The presence of the flagellar maturation process in a metamonad protist Giardia suggests that the basal body or centriole maturation is a universal phenomenon that may represent one of the core processes in a eukaryotic cell.     

Transformation and development of the flagellar apparatus ofCryptomonas ovata (Cryptophyceae) during cell division

Summary InCryptomonas ovata, long, dorsal flagella are produced which transform during the following cell division into short, ventral flagella. At division there is a reorientation in cell polarity, and the parental basal apparatus, which comprises the basal bodies and associated roots, is distributed to the daughter cells via a complex sequence of events. Flagellar apparatus development includes the transformation of a four-stranded microtubular root into a mature root of different structure and function. Each newly formed basal body nucleates new microtubular roots, but receives a striated fibrous root from a parental basal body. The striated roots are originally produced on the transforming basal body and are transferred to the new basal bodies at each successive division. The development of the asymmetric flagellar apparatus throughout the cell cycle is described.     

The flagellar apparatus structure inMicrospora (Chlorophyceae) confirms a close evolutionary relationship with unicellular green algae

The spatial configuration of the flagellar apparatus of the biflagellate zoospores of the green algal genusMicrospora is reconstructed by serial sectioning analysis using transmission electron microscopy. Along with the unequal length of the flagella, the most remarkable characteristics of the flagellar apparatus are: (1) the subapical emergence of the flagella (especially apparent with scanning electron microscopy); (2) the parallel orientation of the two basal bodies which are interconnected by a prominent one-piece distal connecting fiber; (3) the unique ultrastructure of the distal connecting fiber composed of a central tubular region which is bordered on both sides by a striated zone; (4) the different origin of the d-rootlets from their relative basal bodies; (5) the asymmetry of the papillar region which together with the subapical position of the basal bodies apparently cause the different paths of corresponding rootlets in the zoospore anterior; (6) the presence of single-membered d-rootlets and multi-membered s-rootlets resulting in a 7-1-7-1 cruciate microtubular root system which, through the different rootlet origin, does not exhibit a strict 180° rotational symmetry. It is speculated that the different basal body origin of the d-rootlets is correlated with the subapical implant of flagella. It is further hypothesized that in the course of evolution the ancestors ofMicrospora had a flagellar papilla that has migrated from a strictly apical position towards a subapical position. Simultaneously, ancestral shift of flagella along the apical cell body periphery has taken place as can be concluded from the presence of an upper flagellum overlying a lower flagellum in the flagellar apparatus ofMicrospora. The basic features of the flagellar apparatus of theMicrospora zoospore resemble those of the coccoid green algal generaDictyochloris andBracteacoccus and also those of the flagellate green algal genusHeterochlamydomonas. This strengthens the general supposition thatMicrospora is evolutionarily closely related to taxa which were formerly classified in the traditionalChlorococcales.     

Structure of Monocercomonas sp. as Revealed by Electron Microscopy*

SYNOPSIS. Monocercomonas shares many fine-structural features with all other trichomonads. These include the basic arrangement of the kinetosomes as well as of the recurrent and 3 anterior flagella. The pelta-axostyle complex and the parabasal apparatus, i.e. the Golgi complex and the periodic filaments, also conform to the trichomonad pattern. Of interest with regard to the crucial evolutionary position of Monocercomonas, considered to represent the most primitive trichomonad type, is the fact that it has some structures in common with other Monocercomonadidae and Trichomonadinae and others in common with Devescovinidae and Tritrichomonadinae. Among the former organelles are the marginal lamella and the costal base, and among the latter, the comb-like organelle situated between the infrakinetosomal body and parabasal filament 2 as well as the infrakinetosomal body. No traces of either costa or undulating membrane have been noted, but a complex structure homologous to the marginal lamella of Hypotrichomonas and Trichomonadinae is found underlying the short anteriormost portion of the recurrent flagellum that is attached to the body surface. Observations of sections of selected division stages indicate the potential of parental kinetosomes #1 and #3 to become daughter kinetosome #2.