Reconstruction of the flagellar apparatus and microtubular cytoskeleton inPyramimonas gelidicola (Prasinophyceae,Chlorophyta)

Summary The absolute configuration of the flagellar apparatus inPyramimonas gelidicola McFadden et al. has been determined and shows identity withP. obovata, indicating that they are closely related. Comparison with the flagellar apparatus of quadriflagellate zoospores from the more advancedChlorophyceae suggest thatPyramimonas may be a primitive ancestral form. The microtubular cytoskeleton has been examined in detail and is shown to be unusual in that it does not attach to the flagellar apparatus. Cytoskeletal microtubules are nucleated individually, and this is interpreted as an adaptation to the methods of mitosis and scale deployment. In view of the primitive nature of these processes, it is proposed that this type of cytoskeletal organization may represent a less advanced condition than that of the flagellar root MTOCs (microtubule organizing centers) observed in theChlorophyceae.     

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     

Microtubules and the flagellar apparatus in zoospores and cysts of the fungusPhytophthora cinnamomi

Summary A correlated immunofluorescence and ultrastructural study of the microtubular cytoskeleton has been made in zoospores and young cysts ofPhytophthora cinnamomi. Labelling of microtubules using antibodies directed towards tubulin has revealed new details of the arrangement of the flagellar rootlets in these cells, and of the variability that occurs from cell to cell. Most of the variation exists at the distal ends of the rootlets, and may be correlated with differences in cell shape in these regions. The rootlets have the same right and left configuration in all zoospores. The arrangement of the rootlet microtubules at the anterior end of the zoospores raises the possibility that the microtubules on the left hand side of the groove may not comprise an independent rootlet which arises at the basal bodies.The absolute configuration of the flagellar apparatus has been determined from ultrastructural observations of serial sections. In the vicinity of the basal bodies, there is little, if any, variation between individuals, and the structure of the flagellar apparatus is similar to that described for related species of fungi. Two ribbon-like coils surround the central pair of microtubules at the distal tip of the whiplash flagellum, and clusters of intramembranous particles, similar to ciliary plaques, have been found at the bases of both flagella. There are two arrays of microtubules associated with the nucleus in the zoospores. One array lies next to the outer surface of the nuclear envelope, and probably functions in the shaping and positioning of the apex of the nucleus. The nuclear pores in this region are aligned in rows alongside these microtubules. The second array is formed by kinetochore microtubules which extend into a collar-like arrangement of chromatin material around the narrow end of the (interphase) nucleus. During encystment, all flagellar rootlets are internalized when the flagella are detached at the terminal plate. The rootlets arrays are no longer recognizable 5–10 minutes after the commencement of encystment.     

The flagellar root system of zoospores of the green algaChlorosarcinopsis (Chlorosarcinales) as compared withChlamydomonas (Volvocales)

The flagellar root system of zoospores in two species ofChlorosarcinopsis (C. minuta andC. spec.) has been studied in detail. The biflagellate zoospores show a cruciate root system, two of the four microtubular roots containing two microtubules, the other two four microtubules. The flagellar apparatus is otherwise identical with that ofChlamydomonas reinhardi as described byRingo (1967). Evidence is presented that the genusChlamydomonas is characterized by a bilateral symmetric root system (4-2-4-2) rather than a system with four equally numbered roots (i.e. 4-4-4-4). It is suggested that a root system with four identical cruciate roots is not present in any biflagellate algal cell. The taxonomic significance of cruciate root systems in green algae is discussed refering to the identical root systems ofChlorosarcinopsis andChlamydomonas.     

FINE STRUCTURE OF ZOOSPOROGENESIS, ZOOSPORE GERMINATION, AND EARLY GAMETOPHYTE DEVELOPMENT IN CLADOPHORA SURERA (CLADOPHORALES, CHLOROPHYTA)

The fine structure of zoosporogenesis, zoospore germination, and early gametophyte development in Cladophora surera Parodi et Cáceres were studied. Zoosporogenesis started with simultaneous meiosis in all nuclei of apical initial cells. The resulting haploid nuclei duplicated in turn by successive centric, closed mitoses. Then, each initial cell divided into two short zoosporangia. Numerous vacuoles appeared around each sporic nucleus. The delimitation of uninucleate zoosporocytes occurred by cytokinetic furrows produced by the coalescence of tiny, clear vesicles, without microtubules. Final shape of the zoospore resulted from gradual expulsion of vacuoles from the cell body. Mature biflagellate zoospores exhibited a conspicuous apical papilla containing fine granular globules, the basal apparatus, and a microtubular "umbrella" formed by numerous cortical microtubules that ran backward the length of the cell body. The chloroplast showed a conspicuous eyespot. The zoosporangial wall disorganized at the pore through which the zoospores were liberated. Zoospores settled on a substrate by their anterior papilla secreting an adhesive. Germination involved retraction of the apical papilla, loss of the "umbrella" microtubules and eyespot, and the lateral absorption of the entire flagellar apparatus, i.e. basal apparatus plus axoneme, into the cytoplasm. Early gametophyte development involved the synthesis of a thin, young cell wall, the development of outer peripheral vacuoles, the appearance of the marginal reticulate chloroplast, and the formation of the first central vacuoles derived from abundant endoplasmic reticulum. Close to the plasmalemma ran longitudinally oriented cortical microtubules. Eventually, the germling developed an achlorophylic, elongated rhizoidal portion.     

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.     

Surface changes inPhytophthora palmivora zoospores following induced differentiation

Summary Populations ofPhytophthora palmivora zoospores induced to undergo synchronous differentiation by the addition of pectin showed an ordered sequence of surface changes. Within 10 seconds blebs and microvilli appeared on the cell and flagellar surfaces. These projections gave the zoospore a highly irregular and even convoluted surface for up to 30 seconds poststimulation. Between 30 and 40 seconds, pits appeared on the surface, which began to assume a smoother texture. Zoospores then rounded up and by 4 minutes had assumed an almost spherical form. At this time the surface became wrinkled as though shrinkage had occurred. Germ tubes could be seen emerging from almost all cells by 40 minutes, and as germ tubes enlarged, the cell surface became smooth again but with a fibrous texture. Many zoospores retained their flagella through to the germling stage.When cells were stimulated in iso-osmotic rather than hypo-osmotic media, germ tubes in most cases emerged at or very near the site of flagellar attachment. Differentiation in iso-osmotic media showed the same sequence except that the cells did not assume the spherical shape normally characteristic of cysts and germlings differentiating under hypo-osmotic conditions.Strontium-induced differentiation again followed the same general sequence as desribed above, although the cell surface was noticeably more convoluted and irregular prior to the stage of the formation of pits, and flagella were always shed between 40 seconds and 1 minute. Calcium addition up to 20 seconds after the initiation of differentiation with pectin prevented a high proportion of zoospores from encysting, the cells remained motile, and although the frequency of surface blebs and microvilli increased, pits did not appear.These surface changes are consistent with those expected during stimulus-mediated secretion, and it appears that calcium is able to inhibit differentiation only before the secretory step (pit formation).     

The flagellar apparatus and striated rhizoplast of the zoospore ofOlpidium brassicae

Summary The flagellar apparatus and its associated structures of the zoospore ofOlpidium brassicae are described and compared with observations of other zoospores of the uniflagellatePhycomycetes. The zoospore ofO. brassicae is shown to have an extensive cone-shaped rhizoplast fused to both the functional and the vestigial kinetosomes. Three-dimensional reconstructions were made of the kinetosomal region. The vestigial kinetosome differs from the functional, as it only has triplet bundles of microtubules and it lacks a system of props. The proximal termination of the central pair of flagellar microtubules occurs within the axoneme. No terminal plate is observed. The occurrence of dictyosomes in theChytridiales, Monoblepharidales, andHyphochytriales is discussed and it is concluded that a dictyosome may be present in the encysting zoospore and the maturing zoosporangium ofO. brassicae but only vestiges of a dictyosome are to be found in the free-swimming zoospore.     

Microtubules in statocytes from roots of cress (Lepidium sativum L.)

Summary Statocytes in root caps ofLepidium sativum L. were examined by means of ultrathin serial sections to evaluate the amount and distribution of cortical microtubules. The microtubules encircle the cell, oriented normal to the root length axis. In the distal cell edges, microtubules form a network, separating the distal complex of endoplasmic reticulum from the plasmalemma. Preprophase bands in meristem cells are observable rarely, structures which can be regarded as nucleating sites for microtubules are lacking. During ageing of the root cap cells, the number of microtubules increases in combination with a decrease of microtubule length. Development of the roots on a horizontal clinostat preserves a younger developmental stage of the microtubule system regarding amount and length of the individual microtubules. Evidence for an involvement of microtubules in graviperception is low, whereas their role in orienting cellulose microfibrils cannot be ruled out. Compression of the distal network of microtubules after centrifugation of the roots indicates that microtubules in statocytes ofLepidium sativum L. roots might function in stabilizing the distal complex of endoplasmic reticulum.     

Confocal laser scanning microscopy of microtubule organizational changes in isolated generative cells of Allemanda neriifolia during mitosis

Summary Organizational changes in the microtubules of isolated generative cells of Allemanda neriifolia during mitosis were examined using anti--tubulin and confocal laser scanning microscopy. Due to an improved resolution and a lack of out-of-focus interference, the images of the mitotic cytoskeleton obtained using the confocal microscope are much clearer than those obtained using the non-confocal fluorescence systems. In the confocal microscope one can see clearly that the spindle-shaped interphase cells contain a cage-like cytoskeleton consisting of numerous longitudinally oriented microtubule bundles and some associated smaller bundles. At prophase, the shape of the cells invariably becomes spherical. The microtubule cytoskeleton inside the cells concomitantly changes into a less organized form — consisting of thick bundles, patches, and dots. This structural form is not very stable, and soon afterwards the cytoskeleton changes into a reticulate network. Then the nuclear envelope breaks down, and the microtubules become randomly dispersed throughout the cell. Afterwards, the microtubules reorganize themselves into a number of half-spindle-like structures, each possessing a microtubule-nucleating center. The locations of these centres mark out the positions of the presumptive spindle poles. Numerous microtubules radiate from these centres toward the opposite pole. At metaphase, the microtubules form a number of bipolar spindles. Each spindle has two half-spindles, and each half-spindle has a sharply focused microtubule centre at the pole region. From the centres, kinetochore and non-kinetochore microtubules radiate toward the opposite half-spindle. At anaphase A, sister chromatids separate, the cells elongate, and the kinetochore microtubules disappear; the non-kinetochore microtubules, however, remain, and a new array of microtubules, in the form of a cage, appears. The peripheral cage bundles and the non-kinetochore bundles coverge into a sharp point at the pole region. Later, at anaphase B the microtubule cytoskeleton undergoes reorganization giving rise to a new array of longitudinally oriented microtubule bundles in the cell centre and a cage-like cytoskeleton in the periphery. At telophase, some of the cells elongate further, but some become spherical. The microtubules in the central region of the elongated cell become partially disrupted due to the formation of a phragmoplast-junction-like structure in the mid-interzone region. The microtubule bundles at the periphery are spirally organized, and they appear not to be disrupted by the phragmoplast-like junction. The microtubules in the spherical telophase cells (unlike those seen in the elongated telophase cells) are arranged differently, and no phragmoplast-junction-like structure forms in the spherical cells. The structural and functional significances of some of these new features of the organization of the microtubule cytoskeleton as revealed by the confocal microscope are discussed.     

Centrin association with the flagellar apparatus in spores ofPhytophthora cinnamomi

Summary Antibodies raised against the calcium-binding protein centrin, were used to identify and localise centrin containing structures in the flagellar apparatus of zoospores and cysts of the oomycetePhytophthora cinnamomi. Immunoblotting of extracts from zoospores indicates that theP. cinnamomi centrin homologue is a 20 kDa protein. Immunofluorescence microscopy with anti-centrin antibodies reveals labelling in the flagella, the basal body connector and co-localisation along the microtubular R1 root (formerly called AR3) that runs from the right side of the basal body of the anterior flagellum into the anterior of the zoospore close to the ventral surface. The centrin (R1^cen) and tubulin components of the R1 root split into four loops on the right hand side of the ventral groove and rejoin along the left hand side of the groove. The R1 root continues down the left hand side of the zoospore past the basal bodies and parallel to the R4 root. We propose that at least inP. cinnamomi there is no R2 root. Immunogold labelling confirms that centrin is a component of the basal body connector complex. When the zoospores become spherical during encystment, the R1^cen pivots by approximately 90 ° with respect to the nucleus.     

Studies on marine algae of the British Isles. 8. Cystoseira tamariscifolia (hudson) papenfuss

The fine structure of the zoospores of Urospora penicilliformis (Roth) Aresch. (Chlorophyceae) is described. Of special interest is the flagellar apparatus. The proximal part of each of the 4 flagella is ribbon-shaped and contains nine wings attached to the peripheral double tubules. The flagellar root system originates from the flagellar bases and includes striated fibrous roots, passing close to the nucleus, and cruciate nine-stranded microtubular roots along the four corners of the cell. The Golgi bodies produce numerous vesicles, concentrating apically in the cell; they are presumed to be of importance for the attachment of the zoospore.     

The absolute configuration of the flagellar apparatus in zoospores from two species ofLaminariales (Phaeophyceae

Summary We examined the zoospores produced by the unilocular sporangia ofLaminaria digitata (L.) Lamour. andNereocystis luetkeana Post. & Rupr. by serial sectioning to determine the absolute configuration of their flagellar apparatuses. The basal bodies, which are interconnected by three striated bands, lie parallel to the ventral face of the zoospore, and the posterior basal body always is found to the right of the anterior basal body when the cell is viewed from the ventral face, anterior end up. The four rootlets associated with the basal bodies include a major anterior rootlet of about seven microtubules extending from the anterior basal body along the ventral face towards the apex, a five-membered bypassing rootlet that passes ventral to the basal bodies and is connected to the posterior basal body by a posterior fibrous band, and two short rootlets having a single member each, the minor anterior and posterior rootlets. We consider the configuration observed here to be typical of most phaeophycean motile cells. The flagellar apparatus features suggest a considerable phylogenetic difference between thePhaeophyceae and other classes of chlorophyll c-containing organisms.     

Pole reversal and the development of cell asymmetry during division in cryptomonad flagellates

Summary A unique form of cell division is reported for the cellsKomma caudata andCryptomonas ovata (Cryptophyceae). During cytokinesis, the posterior tail-like region of each daughter cell develops from the anterior region of the parental cell. This process, termed pole reversal, involves a major realignment in overall cell polarity as well as alterations to cytoplasmic and surface components. Pole reversal may be a consequence of flagellar apparatus transformation and reorientation during division, and pole reversal may facilitate the development of the asymmetric cell shape in daughter cells.     

Effect of temperature,hydrogen ion concentration and osmotic potential on zoospore production by threePythium species isolated from pond water

Pythium fluminum produced zoospores most abundantly at 15°C, whereas the optima forPythium group F andP. marsipium were 20 and 25°C, respectively. Increasing the incubation temperature above the optimum resulted in the decrease of the duration of zoospore production. InPythium group F the ability to produce zoospores was not lost even after incubation at 40°C for 24 h. On the other hand,P. marsipium andP. fluminum lost the ability under these conditions. Zoospore production was inhibited at pH 4.5 and 10.5 in all the species tested.Pythium fluminum andP. marsipium were found to have two pH optima for zoospore production (7.5 and 9.5 for the former and 5.5 and 8.5 for the latter). The optimum pH for zoospore production byPythium group F was 6.5–7.5. Moderate osmotic potentials (–0.27–0.47 MPa) appeared to favor zoospore production by the pythia tested. The effect of temperature, pH and osmotic potential on zoospore production was discussed in relation to pollution of pond water.     

Reproduction by zoospores inOedogonium

Summary Emergence of zoospores ofOedogonium and their subsequent developmental changes have been studied using live material and sections prepared for light and electron microscopy. Release commences with rupture of the cell wall at its pre-weakened site near the apical caps. The pliable protoplast of the zoospore becomes completely spherical once free of the wall; it is enclosed within the hyaline vesicle which expands continuously and then disappears. Meanwhile, as the flagella become active, the zoospore begins to elongate and its dome starts to protrude from a circular constriction where the flagella are inserted. Once free of the hyaline vesicle, it is actively motile for a variable period, during which elongation continues. The motile phase ceases when the zoospore begins to vibrate, whereupon the flagella are all violently shed. Soon after this, the constriction disappears from around the dome which becomes more pointed; the immobile cell now elongates further, increasing in volume. The cell periphery contains numerous contractile vacuoles. Zoospore elongation may be associated with a proliferation of longitudinal microtubules, and once the flagella are shed, the flagellar rootlet system disintegrates, probably releasing the rootlet microtubules. Mechanisms involved in the release of the zoospore are also discussed.     

Flagellar apparatus ultrastructure inMesostigma viride (Prasinophyceae)

The ultrastructure of the flagellar apparatus ofMesostigma viride Lauterborn (Prasinophyceae) has been studied in detail with particular reference to absolute configurations, numbering of basal bodies, basal body triplets and flagellar roots. The two basal bodies are interconnected by three connecting fibers (one distal fiber = synistosome, and two proximal fibers). The flagellar apparatus shows 180° rotational symmetry; four microtubular flagellar roots and two system II fibers are present. The microtubular roots represent a 4-6-4-6-system. The left roots (1s, 2s) consist of 4 microtubules, each with the usual 3 over 1 root tubule pattern. Each right root (1d, 2d) is proximally associated with a small, but typical multi-layered structure (MLS). The latter displays several layers corresponding to the S₁ (the spline microtubules: 5–7), and presumably the S₂—S₄ (the lamellate layers) of the MLS of theCharophyceae. At its proximal origin (near the basal bodies) each right root originates with only two microtubules, the other spline microtubules being added more distally. The structural and positional information obtained in this study strongly suggest that one of the right roots (1d) ofMesostigma is homologous to the MLS-root of theCharophyceae and sperm cells of archegoniate land plants. Thus the typical cruciate flagellar root system of the green algae and the unilateral flagellar root system of theCharophyceae and archegoniates share a common ancestry. Some functional and phylogenetic aspects of MLS-roots are discussed.Dedicated to Prof. DrLothar Geitler on the occasion of his 90th birthday.     

Thrust reversal by tubular mastigonemes: immunological evidence for a role of mastigonemes in forward motion of zoospores ofPhytophthora cinnamomi

Summary The role of tubular mastigonemes in the reversal of thrust of the anterior flagellum ofPhytophthora cinnamomi was analysed using mastigoneme-specific monoclonal antibodies and immunoflu-orescence and video microscopy. Exposure of live zoospores ofP. cinnamomi to the mastigoneme-specific Zg antibodies caused alterations in the arrangement of mastigonemes on the flagellar surface and at Zg concentrations above 0.3 /ml, mastigonemes became detached from the flagellum. As a consequence of antibody binding to the mastigonemes there were concentration-dependent perturbations in zoospore swimming behaviour and anterior flagellum beat pattern. With increasing antibody concentration zoospores swam more slowly and other parameters of their swimming pattern, such as the wavelength of the swimming helix and the frequency of rotation, were also reduced. The effects of Zg antibodies were specific at two levels: control immunoglobulins or antibodies that bound to other flagellar surface components did not have an effect on motility, and Zg antibodies did not interfere with the motility of zoospores of oomycete species to which they did not bind. The effects of antibody-induced disruption of mastigoneme arrangement strongly support previous hypotheses that tubular mastigonemes are responsible for thrust reversal by the anterior flagellum, enabling it to pull the cell through the surrounding medium.     

Outer segment growth and periciliary vesicle turnover in developing photoreceptors of Xenopus laevis

Summary Cytoskeletal alterations in the cytoplasm of chromatolytic neurons of the dorsal root ganglia were studied in chickens after transection of the sciatic nerves. These studies were carried out using cryofixation with a nitrogencooled propane jet. By this method, the morphological complexity of the cytoskeleton in normal perikarya and cell processes can be visualized. The cytoskeleton of the dorsal root ganglion cells (DRG) is composed of an intricate network of microtubules, neurofilaments and microfilaments. The membrane-bounded cell organelles, as well as the cell nucleus and the plasmalemma, are linked to the microtubules and neurofilaments by microfilaments (or crosslinkers). As a result of the transection of the axon, chromatolysis takes place, characterized by dislocation of cell organelles, eccentric position of the nucleus and dispersion of the parallel cisternae of the rough endoplasmic reticulum throughout the cytoplasm. This characteristic phenomenon coincides with a regression of the neurocytoskeletal network. The neurofilaments and microtubules become shorter, and the microfilaments are replaced by strands of globular or granular material. The temporary regression of the microfilaments leads to a dispersion of the cell organelles. During the remodelling of the cytoskeletal structures, proliferation of the neurofilaments in the regenerating neurons may occasionally be observed. These results show that the cytoskeletal structures are responsible not only for the preservation of cell shape, but also for the maintenance of the normal distributional pattern (location and mobility) of the intracellular components.