THE FINE STRUCTURE OF MITOSIS AND CELL DIVISION IN THE CHRYSOPHYCEAN ALGA OCHROMONAS DANICA1

At the ultrastructural level, cell division in Ochromonas danica exhibits several unusual features. During interphase, the basal bodies of the 2 flagella replicate and the chloroplast divides by constriction between its 2 lobes. The rhizoplast, which is a fibrous striated root attached to the basal body of the long flagellum, extends under the Golgi body to the surface of the nucleus in interphase cells. During proprophase, the Golgi body replicates, apparently by division, and a daughter rhizoplast, appears. During prophase, the 2 pairs of flagellar basal bodies, each with their accompanying rhizoplast and Golgi body, begin to separate. Three or 4 flagella are already present at this stage. At the same time, there is a proliferation of microtubules outside the nuclear envelope. Gaps then appear in the nuclear envelope, admitting the microtubules into the nucleus, where they form a spindle. A unique feature of mitosis in O. danica is that the 2 rhizoplasts form the poles of the spindle, spindle microtubules inserting directly onto the rhizoplasts. Some of the spindle microtubules extend from pole to pole; others appear to attach to the chromosomes. Kinetochores, however, are not present. The nuclear envelope breaks down, except, in the regions adjacent, to the chloroplasts; chloroplast ER remains intact throughout mitosis. At late anaphase the chromosomes come to lie against part of the chloroplast ER. This segment of the chloroplast ER appears to be incorporated as part of the reforming nuclear envelope, thus reestablishing the characteristic nuclear envelope—chloroplast ER association of the interphase cell.     

Ultrastructure of the biflagellate gametes of Collinsiella cava (Ulvophyceae, Chlorophyta)

The fine structure of the biflagellate gametes of Collinsiella cava (Yendo) Printz was investigated in detail to clarify the species's taxonomic and phylo‐genetic position. Gametes are covered by small square scales with no distinct substructure. The chloroplast of the gamete includes an eyespot comprised of two layers of globules, and a pyrenoid that is traversed by one or a few thylakoids. Basal bodies overlap at their proximal ends and are offset in a counterclockwise orientation. Each basal body has a small bipartite terminal cap, a prominent proximal sheath comprised of two unequal subunits and a circular element situated at the cartwheel portion. A distal fibre, a connecting fibre and linkage between proximal sheaths connect the two basal bodies. Microtubular roots are comprised of two dexter (d) roots, subtended by the system I fibre, and two sinister (s) roots. Gametes have a single rhizoplast which extends parallel to one of the two d roots and extends to the mating structure. The ultrastructure of Collinsiella gametes is very similar to that of Mono‐stroma and other members of the Ulotrichales, Ulvophyceae, and we concluded that the genus Collinsiella should be treated as a member of the Monostromat‐aceae. The planozygote has four basal bodies, eight microtubular roots and two eyespots always situated at the same face of the cell. From observations of the planozygotes, the position of the mating structure relative to the flagellar apparatus is not consistent, but converse, between two mating types. A comparison of the location of the mating structure in Chlamydomonas and other green algae is presented.     

Ultrastructural and immunocytological studies on the rhizoplast in the chrysophycean alga Ochromonas danica

The rhizoplast, a striated band elongating from the flagellar basal body to the nucleus, is conspicuous in cells of Ochromonas danica Prings. In interphase cells, it runs from the basal body of the anterior flagellum to the space between the nucleus and the Golgi body. In O. danica, the rhizoplast duplicates during mitosis and the two rhizoplasts serve as mitotic poles. In the present study, we reinvestigated mitosis of O. danica using transmission electron microscopy and immunofluorescence microscopy, especially focusing on the rhizoplast. The nuclear envelope became dispersed during metaphase, and the rhizoplasts from two sets of the flagellar basal bodies functioned as the mitotic poles. Immunofluorescence microscopy using anti‐α‐tubulin, anti‐centrin and anti‐γ‐tubulin antibodies showed that centrin molecules were localized at the flagellar basal bodies, whereas γ‐tubulin molecules were detected at the rhizoplast during the whole cell cycle.     

The flagellar root apparatus,the microtubular system and associated organelles in the chrysophycean flagellate,Poterioochromonas malhamensis Peterfi (syn.Poteriochromonas stipitata Scherffel andOchromonas malhamensis pringsheim)

Summary The two flagella ofPoterioochromonas are inserted in an apical platform which is shaped by six long flagellar root fibres. The arrangement and structure of these root fibres are described in detail. One of these fibres is the single nucleating site for cytoplasmic interphase microtubules which extend peripherally down to the cytoplasmic tail. Another fibre proceeds toward the centre of the cell and passes the nucleus but is different in structure, position and function from the striated rhizoplast found in many chrysophycean flagellates which is observed but vestigial inPoterioochromonas.A specific kinetosomal mitochondrion has a threefold attachment to the flagellar root apparatus. The chloroplast is also bound to the root system. It has no stigma, but a special continuation of the periplastidial cisterna is developed instead. Another cisterna extends from the nuclear envelope-dictyosome interspace to the kinetosome of the long flagellum. The functional and taxonomic meanings of these structures and of their mutual arrangement are discussed. It is concluded that the present strain (no. 933-1 a of the Collection of Algal Cultures at the Institute of Plant Physiology, Göttingen) has to be excluded from the genusOchromonas.     

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.     

Fine structure of photo-kinetic systems in Dinobryon cylindricum var. Alpinum (chrysophyceae)

The ultrastructure of the stigma and associated flagellar-microtubular systems in Dinobryon cylindricum var. alpinum is described in detail and compared with observations on comparable photo-kinetic systems in other chrysophycean organisms. The chloroplastidic stigma of D. cylindricum var. alpinum is shown to lie in a particular positional relationship to the flagellar swelling in the anterior furrow and to several other organelles, to consist of a monolayer of c. 40 pigmented granules, each c. 250–500 nm diameter, arranged in a definite pattern, and to be overlain by several membrane systems. Other cytoplasmic pigmented bodies with dense crystalline contents surrounded by a single “unit membrane” aggregate near the anterior furrow on the side opposite the stigma. The swelling on the proximal portion of the smooth flagellum is separated from the plasmalemma of the anterior furrow by a nearly constant distance of 75–100 nm, has a multilamellate substructure that is linked by fine radiating interconnections to the axoneme doublets, and is connected to the plasmalemma by a system of fibrillar interconnections. A transitional helix in the basal body region is described as similar to structures reported in other chrysophycean flagellates. A striated rhizoplast with a periodicity of c. 90 nm extends from basal body I to the nuclear envelope. A seven-stranded microtubular root extends from the same basal body. Other fibrous and microtubular root systems are also described. The inter-relationships and possible functions of the aforementioned structures are discussed.     

Analysis of cytoplasmic microtubules and flagellar roots in the zoospores ofAllomyces macrogynus

Summary Cytoskeletal and flagellar microtubules in the zoospores of the aquatic fungusAllomyces macrogynus are resistant to microtubule depolymerizing drugs. Consequently, we have analyzed the partial composition and organization of microtubules (Mts) in the cytoplasm and flagellar apparatus in the zoospores ofA. macrogynus. Evidence from two-dimensional gel electrophoresis demonstrated the presence of two -tubulin isoforms in axonemal and cytoplasmic Mts. In addition, a monoclonal antibody specific for acetylated -tubulin was used on one-dimensional protein blots to show that acetylated -tubulins are present in isolated zoospore cell bodies and axonemes. Immunofluorescence microscopy observations using this monoclonal antibody demonstrated that flagellar, kinetosomal, and cytoplasmic Mts were labeled. The nature of Mts in the flagellar apparatus was studied ultrastructurally. InA. macrogynus, the flagellar apparatus consists of the kinetosome, rhizopolast (striated flagellar rootlet), axoneme, and 9 sets of triplet Mts which radiate anteriorly from the proximal end of the kinetosome (microtubular rootlet), Analysis of the rhizoplast indicated that this structure does not contain Mts. The rhizoplast, which connects the functional kinetosome with a single, large basal mitochrondrion, consists of four electron-opaque bands. Serial-sectioning indicated that the rhizoplast is always adjacent to kinetosome triplets 1, 2, and 9, and thus lies perpendicular to the plane of flagellar beat. These results suggest that the primary function of the rhizoplast is to organize the kinetosome and mitochondrion with respect to one another and to bias flagellar beat in the appropriate orientation for cell motility.Abbreviations BSA bovine serum albumin - BCA bicinchoninic acid - DS dilute salts - EGTA ethylene glycol-bis-(-aminoethyl ether)-N,N-tetracetic acid - EM electron microscopy - Mes 2-(N-morpholinomethane sulfonic acid - Mt microtubule - NP-40 Nonidet P-40 - 1-D PAGE one-dimensional polyacrylamide gel electrophoresis - PBS phosphate-buffered saline - PMSF phenylmethylsulfonyl fluoride - SDS sodium dodecyl sulfate - 2-D PAGE two-dimensional polyacrylamide gel electrophoresis - Tween-20 polyoxyethylenesorbitan monolaurate     

Der Rhizoplast vonTetraselmis cordiformis sichtbar auch im Lichtmikroskop

Summary The massive rhizoplast of the prasinophycean algaTetraselmis cordiformis, well known from the electron microscopy investigations, is shown on living cells by light microscopy using the phase-contrast. It appears as a dark girdle attached to the surface of the nuclear envelope.

     

The main cell morphology of the freshwater colorless chrysomonad Spumella sp. (Ochromonadales, Chrysophyceae)

The cell structure of the freshwater chrysomonad Spumella sp. has been studied. The cell contains a vesicular nucleus, mitochondria with tubular cristae, Golgi apparatus, flagellar roots, and wide dorsal microtubular band. The flagella bear the spiral of four to five coils in the transitional zone. The rudiments of mastigonemes have been found in the perinuclear space. The compact leucoplast has an amorphous core surrounded by the membrane. No stigma has been detected. The leucosin vacuole, rhizoplast, and swelling of the short flagellum are absent. One to three osmiophilic granules lie near the leucoplast. The contractile vacuole is surrounded by tubules. The resemblance and difference of investigated flagellate with other chrysomonads are discussed.     

Elektronenmikroskopische Untersuchung über die Vorbereitung der Infektion in encystierten Zoosporen von Olpidium brassicae

Zusammenfassung Anhand der Ultrastrukturen wird die präinfektionelle Phase an die Wirtszelle angehefteter Olpidium-Zoosporen beschrieben. Vor der Encystierung wird die Geißel eingezogen; sie liegt dann in zwei Windungen in der Zoospore. Beim Abbau des Geißelapparates schiebt sich als erstes das Dictyosom der Zelle zwischen den Rhizoplasten und das Kinetosom sowie das Centriol. Nach Auflösung des Rhizoplasten trennt sich das Axonema vom Kinetosom und desintegriert. Die Centriolen lagern sich schließlich zwischen Kern und Dictyosom. Vom Dictyosom produzierte Sekretvesikeln sammeln sich an der Anheftungsstelle in der Cyste; dort organisieren sich oft auch Lomasomen-ähnliche Strukturen. Im distalen Teil der Zelle entsteht aus Elementen des ER über ein multivesiculäres Stadium die Vacuole.

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Electron microscopical study on the preparation of infection in encysted zoospores of Olpidium brassicae

Summary The ultrastructure of the preinfectional phase of zoospores of Olpidium brassicae attached to host cells is described. Prior to encystment the flagellum is retracted in the zoospore, where it forms two windings. At the beginning of flagellum desintegration the dictyosome is going to separate the rhizoplast from the kinetosome and the centriole. After desintegration of the rhizoplast the axonema is separated from the kinetosome and desintegrates too. The resulting two centrioles are finally located between the nucleus and the dictyosome. Vesicles produced by the dictyosome are aggregating at the region of attachment of the cyste; there often are found lomasome-like structures. The vacuole is formed in the distal region of the cyst from an multivesicular structure originating from cisternae of the ER.

     

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     

ZOOSPORE STRUCTURE OF THE MYCOPARASITIC CHYTRID CAULOCHYTRIUM PROTOSTELIOIDES OLIVE

The subcellular organization of zoospores released from sessile, parasitic sporangia of Caulochytrium protostelioides was studied with light and electron microscopy. A single flagellum is posteriorly directed but laterally inserted into the cylindrical motile zoospore. A striated rhizoplast attaches the proximal end of the kinetosome to a specialized region of the nuclear envelope. A system of rough endoplasmic reticulum, smooth endoplasmic reticulum, dictyosomes and bristle-coated vesicles are associated with the one to several pulsating vacuoles typically located near the flagellar apparatus. The microbody-lipid globule complex (MLC) comprises one to many lipid globules. An extensive microbody branches around each lipid globule and encloses a portion of the rhizoplast. A reticulum of smooth surfaced cisternae interdigitates among the branches of the complex microbody, and cisternae are opposed to the surface of lipid globules opposite the microbodies. Mitochondria with predominantly circular profiles are scattered throughout the zoospore body, but several are always adjacent to the microbody, and hence, are also part of the MLC. Ribosomes are uniformly distributed throughout the zoospore, and one to several cisternae of rough endoplasmic reticulum are adjacent to the nuclear envelope. Zoospores of C. protostelioides are similar to several other chytrid zoospores, which also have the same type of microbody-lipid globule complex, but yet are structurally distinct from any other chytrid zoospore.     

MICROTUBULE BIOGENESIS AND CELL SHAPE IN OCHROMONAS : I. The Distribution of Cytoplasmic and Mitotic Microtubules

In the first of two companion papers which attempt to correlate microtubules and their nucleating sites with developmental and cell division patterns in the unicellular flagellate, Ochromonas, the distribution of cytoplasmic and mitotic microtubules and various kinetosome-related fibers are detailed. Of the five kinetosome-related fibers, which have been found in Ochromonas, two, the kineto-beak fibers and the rhizoplast fibers are utilized as attachment sites for distinct groups of microtubules. The set of microtubules attached to the kineto-beak fibers apparently shape the anterior beak region of the cell whereas the rhizoplast microtubules appear to extend into and shape the tail in vegetative cells. In mitotic cells a rhizoplast is found at each spindle pole apparently serving as foci for the spindle microtubules. These findings are discussed in relation to the less well defined attachment sites for vegetative and mitotic microtubules in other kinds of cells. It is noted that the effects of depolymerizing microtubules in vivo might be easily quantitated in whole populations since no external wall or pellicle contributes to the maintenance or the biogenesis of the characteristic cell form of Ochromonas.     

Absolute orientation of the flagellar apparatus of Hibberdia magna comb. nov. (Chrysophyceae)

The first flagellum of Hibberdia magna comb. nov. bears mastigonemes that have both short and long lateral filaments attached to the tubular shaft. The second flagellum is very short (ca. 850 nm) and is directed posteriorly approximately 160° from the first flagellum. Three microtubular flagellar roots (R₁, R₂ and R₄) and a rhizoplast (= striated root) are present. The R₁ root consists of four microtubules that arise near the right surface of the first flagellum basal body; the R₁ root extends to the dorsal side of the cell and then curves back along the left side of the cell. Cytoskeletal microtubules are nucleated from the R₁ root including one loose cluster of cytoskeletal microtubules that extends down the left side of the cell adjacent to the contractile vacuole. The R₂ root is a single microtubule that arises along the left surface of the first flagellum basal body and extends to the left side of the cell. The R₄ root consists of three microtubules that arise along the left side of the second flagellum basal body. A helical band wraps around two microtubules at the proximal end of the R₄ root. Two of the three R₄ root microtubules extend along the left side of the second flagellum, curve around to the right side of that flagellum and terminate. No R₃ root was found. The orientation of the basal bodies of Hibberdia gen. nov. is similar to that of the Xanthophyceae and Oomycetes. There are apparent homologies in the R₁, R₂ and R₄ roots of Hibberdia and these and other protists, but only Hibberdia lacks a R₃ root. Three long flagella are present in preprophase but later one is endocytosized and the axoneme extends to the posterior of the cell. During metaphase the nuclear envelope is more or less intact except at the poles; the flagellar apparatuses are at the poles and the spindle microtubules originate near the basal bodies. Two stages are known in the life history: 1) a capsoidlike state with non-swimming flagellate cells inside a colonial gel, and 2) a free-swimming single-celled monad state. Vegetative cell division occurs in both stages. The flagellar apparatus, the cell division process and the life history combined with the previously described unique light-harvesting antheraxanthin make H. magna distinct from other algae. A new genus, Hibberdia gen. nov., a new family, Hibberdiaceae fam. nov. and a new order, Hibberdiales, ord. nov. are described.     

Light and electron microscopical studies onHafniomonas gen. nov. (Chlorophyceae,Volvocales), a genus resemblingPyramimonas (Prasinophyceae)

Based on light and electron microscopical studies ofPyramimonas reticulata the genusPyramimonas is shown to contain a number of unrelated flagellates.P. reticulata andP. montana are transferred to the new genusHafniomonas, cells of which differ fromPyramimonas in shape, in the absence of scales and hairs on the body and flagellar surfaces, in details of the chloroplast, the position of the nucleus, the Golgi apparatus, the internal structure of the flagellar apparatus, and in cell division. The prasinophytePyramimonas contains a characteristic association of a large microbody and a rhizoplast, situated on the nuclear surface. A similar association is being found in an increasing number of prasinophycean flagellates, but is absent inHafniomonas, which is considered related to chlorophycean rather than prasinophycean flagellates. The phylogenetic position ofHafniomonas is discussed, based in particular on details of the unique flagellar apparatus.     

THE FLAGELLAR APPARATUS OF CHRYSOLEPIDOMONAS DENDROLEPIDOTA (CHRYSOPHYCEAE), A SINGLE-CELLED MONAD COVED WITH ORGANIC SCALES1

The flagellar apparatus of Chrysolepidomonas dedrolepidota Peters et Andersen is similar to that of other members of the Ochromonadales, Chrysophyceae. there are four microtubular roots (R_1-4) and a system II fiber (= rhizoplast). the R₁ root consists of three microtubules that nucleate many cytoplasmic microtubules. One compressed band of 10 or more cytoplasmic microtubules is directed black along the R1 root in an anti-parallel direction. The R₂ root consists of one to two microtubules, and it extends toward the distal end of the R₁ root. The R₃ root consists of six (?seven) microtubules near its proximal end. The “a” and “f” microtubules of the R₃ root are under the short flagellum, and the “f” microtubule loops back and under the basal body, extending down to the nucleus. The R₄ root consists of one to two microtubules extending along the left side of the shot flagellum and curving under the short flagellum where it terminates near the “a” microtubule of R₃ Both flagella have a transitional plate and a transitional helix with five gyres. There is a thin, second plate in the basal body at the level of the distal end of the “c” tubules of the basal body triplets. The tripartite flagellar hairs have long lateral filaments but lack short lateral filaments. We compare the flagellar apparatus with that of other members of the Ochromonadales and members of the Hydrurales and Hibberdiales.     

The zoospore ofOlpidium brassicae

Summary The ultrastructure of the zoospore ofOlpidium brassicae is described and compared with observations made of other zoospores of the uniflagellatePhycomycetes. The zoospore ofO. brassicae is characterized by an extensive, cone-shaped rhizoplast and a lack of a nuclear cap, as well as a side-body complex or a rumposome. Vacuoles which contain osmiophilic material are termed gamma-like particles. Three-dimensional reconstructions based on serial sectioning were made of the organelles in the region of the nucleus, showing that the zoospore ofO. brassicae contains one or at most two elaborately branched mitochondria. Microbodies have a high degree of interconnection and are in intimate association with the mitochondrion, lipid drops, and the nuclear envelope.     

An ultrastructural comparison betweenSpermatozopsis andDunaliella (Chlorophyceae)

The ultrastructure of the type species of the genusDunaliella, D. salina, has been reinvestigated in an attempt to clarify the relationships betweenDunaliella andSpermatozopsis. Dunaliella salina differs in the following ultrastructural characters fromSpermatozopsis (as exemplified byS. similis Preisig etMelkonian): presence of a distinctive surface coat covering the plasmalemma; presence of a prominent pyrenoid (with pairs of thylakoids partially entering the pyrenoid matrix); dictyosomes parabasal; endoplasmic reticulum closely underlying the plasmalemma around most of the cell; contractile vacuoles absent; cell form ovoid to elongated and not spirally twisted; mitochondrial profiles near the flagellar apparatus. Differences in the ultrastructure of the flagellar apparatus: basal body angle more or less fixed; distal connecting fibre cross-striated; system II fibre (rhizoplast) present, associated with mitochondrial profile; system I fibre underlying two-stranded microtubular root; mating structure present. These ultrastructural differences justify distinction between the two taxa at generic level. The problematical status of freshwater species ofDunaliella is briefly discussed.     

DEVELOPMENT OF THE FLAGELLAR APPARATUS OF NAEGLERIA

Flagellates of Naegleria gruberi have an interconnected flagellar apparatus consisting of nucleus, rhizoplast and accessory filaments, basal bodies, and flagella. The structures of these components have been found to be similar to those in other flagellates. The development of methods for obtaining the relatively synchronous transformation of populations of Naegleria amebae into flagellates has permitted a study of the development of the flagellar apparatus. No indications of rhizoplast, basal body, or flagellum structures could be detected in amebae. A basal body appears and assumes a position at the cell surface with its filaments perpendicular to the cell membrane. Axoneme filaments extend from the basal body filaments into a progressive evagination of the cell membrane which becomes the flagellum sheath. Continued elongation of the axoneme filaments leads to differentiation of a fully formed flagellum with a typical "9 + 2" organization, within 10 min after the appearance of basal bodies.     

The rhizoplast of chrysomonads, a basal body–nucleus connector that polarises the dividing spindle

Summary. An ultrastructure study of the rhizoplast in Synura petersenii, Mallomonas fastigiata, and M. insignis shows that it consists of 15–20 striated rootlets that form a claw or an incomplete cone over the nucleus. These rootlets course along one face of the nucleus between the nuclear membrane and the cis-face of the Golgi stack of cisternae. They converge and merge above the nucleus, forming a stub attached to the proximal section of the two basal bodies. These cross-striated rootlets are composed of closely packed longitudinal microfibrils. By immunofluorescence, the basal bodies and the rootlets forming the claw were decorated by the anti-centrin monoclonal antibody ICL19 raised against the Paramecium tetraurelia acidic centrin protein and by two antibodies raised against the striated parabasal and costal striated fibres of trichomonads. Only the anti-centrin monoclonal antibody 20H5 raised against Chlamydomonas reinhardtii centrin strongly labelled the 20–22 kDa protein bands from the extracted cytoskeleton of S. petersenii by immunoblotting. Electron micrographs of mitosis in S. petersenii cells revealed that the segregated pairs of basal bodies are linked by the striated rootlets of the rhizoplast to the poles of the mitotic spindle. The spindle microtubules arise perpendicularly from the striated rootlets of the basal body–nucleus connector forming the centrosome. In conclusion, in these cells there is a basal body–nucleus connector similar to that of C. reinhardtii and other chlorophytes. It contains centrin proteins, it is involved in the linkage of the basal bodies to the nucleus and is a component of the spindle pole body or centrosome in the dividing cell.Correspondence and reprints: Biologie des Protistes, Université Blaise Pascal de Clermont-Ferrand, Campus des Cézeaux, 63177 Aubière Cedex, France.Present address: Romagnat, France.Received February 7, 2003; accepted May 21, 2003; published online September 23, 2003