ULTRASTRUCTURAL STUDIES OF SPERMATOGENESIS IN THE ANTHOCEROTALES. I. THE BLEPHAROPLAST AND ANTERIOR MITOCHONDRION IN PHAEOCEROS LAEVIS: EARLY DEVELOPMENT

Electron microscopic examination of thin sections showed that the blepharoplast of a young spermatid of Phaeoceros consists of two side-by-side centrioles and an accumulation of osmiophilic, granular matrix at their proximal ends. Lying between these nearly parallel organelles is a dark-staining body that will later disappear at the onset of flagellogenesis. For a brief period the centrioles are oriented perpendicular to the nuclear surface so that the granular matrix at their proximal ends is confluent with the nuclear envelope; furthermore, the nucleoplasm immediately in front of the centrioles becomes densely staining. The multilayered structure (MLS) develops directly under the centrioles. It comprises a band of 12 microtubules (the S₁ stratum) and three lower strata (S_2–4) whose constitutent lamellae are oriented at an oblique angle to the S₁ axis. While the S₁ tubules grow rearward over the nucleus which forms a beak adjacent to the posterior end of the lamellar strata, the centrioles are transformed into basal bodies with the distal growth of the axonemes and the proximal growth of the central cartwheels and lowermost triplets. The proximal ends of the basal bodies and the S₁ tubules overlying the lamellar strata are invested with osmiophilic matrix that extends down to the S₂ layer and may temporarily occlude the lamellar plates. At the onset of nuclear elongation an anterior mitochondrion becomes situated close beneath the lamellar strata which extend laterally beyond the S₁ tubules.     

SPERMATOGENESIS IN COLEOCHAETE PULVINATA (CHAROPHYCEAE): EARLY BLEPHAROPLAST DEVELOPMENT

Transmission electron microscopy of serial thin sections was used to reconstruct several early developmental stages of the blepharoplast in Coleochaete pulvinata spermatids. These were compared to published studies of blepharoplast development in Charales and the closest relatives of charophycean green algae among embryophytes, i.e., hornworts and liverworts. Bicentriolar centrosomes such as occur in bryophytes and fern allies were not observed in Coleochaete. Centriole replication in C. pulvinata was orthogonal as in Charales. The resulting two daughter centrioles were oriented perpendicularly and joined proximally by electron-dense material. Their orthogonal relationship was maintained throughout blepharoplast development by a massive, banded connective which appeared early. In spermatids of hornworts and liverworts, a multilayered structure (MLS) develops in association with two centrioles destined to become flagellar basal bodies. When the MLS of these lower land plants is sectioned at right angles to the long axis of the microtubular layer, the MLS is observed to lie beneath cross sections of both centrioles. In contrast, when developing MLSs of C. pulvinata and Charales are similarly sectioned, they occur beside a cross section of just one of the two centrioles. In C. pulvinata (as in other charophytes), MLS lamellae are oriented at a 90-degree angle to the long axis of the S₁ microtubules from the beginning. This contrasts with the 40–45 degree angle between the MLS lamellae and S₁ microtubules universally reported for archegoniates. In early C. pulvinata spermatids, spline microtubules are closely associated with an anterior mitochondrion having a low stromal density and few cristae. An anterior mitochondrion is typically associated with blepharoplast development in hornworts and liverworts, but has not previously been reported to occur in Coleochaete or any other charophycean alga. In Coleochaete, as in hornworts and liverworts, but unlike Charales, structure of mature blepharoplasts reflects early blepharoplast ontogeny. Very little change in positional relationships among blepharoplast components (flagella, connective, MLS) occurs during development. These character-state differences are of importance in cladistic analyses of charophycean algae and lower land plants.     

Fine structural studies of the bipolarization of the mitotic apparatus in the fertilized sea urchin egg. I. The structure and behavior of centrosomes before fusion of the pronuclei

During fertilization the sperm brings two centrosomes into the egg. One centrosome contains a centriole of normal length originally seen as the basal body of the sperm flagellum. Characteristically, the proximal half is enwrapped in osmiophilic material. This centrosome is attached to the centrosomal fossa, a bowl-shaped depression of the nuclear envelope of the male pronucleus. Microtubules radiate out from the osmiophilic half characterizing this structure as a centrosome and microtubule organizing center (MTOC). The second centrosome which also acts as an MTOC is attached to the mitochondrion of the sperm. At the beginning it appears as an unstructured accumulation of osmiophilic material out of which later on centriolar microtubules grow. Though this centrosome is marked by an immature centriole it is capable of organizing microtubules and of reproducing itself. This centrosome becomes loosely associated with the female pronucleus by means of microtubules. Then it separates from the mitochondrion which finally is lost. When the two pronuclei fuse, the centrosome derived from the basal body remains firmly attached to the centrosomal fossa, which has persisted in the envelope of the zygote nucleus after pronuclear fusion. Using the fossa as a marker of the position of this centrosome on the nuclear surface, we conclude that it is a stationary centrosome in the process of bipolarization for the first mitosis.     

ULTRASTRUCTURAL STUDIES OF SPERMATOGENESIS IN THE ANTHOCEROTALES. II. THE BLEPHAROPLAST AND ANTERIOR MITOCHONDRION IN PHAEOCEROS LAEVIS: LATER DEVELOPMENT

Chromatin condensation begins as the multilayered structure (MLS) in Phaeoceros reaches its maximally structured differentiation. As nuclear elongation and chromatin condensation proceed, the S_2–4 strata disappear, and the nuclear beak extends between the S₁ and the nearly spherical anterior mitochondrion. In a mature sperm the mitochondrion is elongate and lies completely anterior to the blunt front end of the nucleus. The 12 S₁ tubules extend over the anterior mitochondrion and nucleus, but their number becomes reduced to five at the level where the nucleus' midportion is constricted. The anterior ends of the S₁ tubules lie embedded in a rather conical osmiophilic crest. Flagellar insertion is restricted to the extreme anterior tip of the S₁ tubules. The locomotory apparatus in Phaeoceros is compared with that of other bryophytes.     

Spermatogenesis of a marine snail,Littorina sitkana

Summary The fine structure of the spermatogonium, spermatocyte and spermatid of a marine snail, Littorina sitkana is described. The ring centriole (annulus) is formed from the distal centriole and it migrates to the base of the mitochondrial region where it lies in a joint-like structure which is formed by an area of invaginated plasma membrane. The distal and proximal centrioles are at first perpendicular to each other but the proximal centriole rotates to a position coaxial with the distal centriole and fuses with it. The peripheral doublet fibers are continuous between the two centrioles but the central fibers originate only in the distal centriole. The acrosome differentiates from the proacrosomal granule which is derived from a Golgi body. Microtubules, present at this stage, may assist acrosomal formation. Chromatin condensation begins with the formation of fibrous strands, then to lamellar plates which become folded and later twisted around the flagellar shaft. In the final stages the lamellae appear in cross section as concentric rings which eventually fuse to form a homogeneously dense nuclear tube.     

LIGHT MICROSCOPE AND ULTRASTRUCTURAL STUDIES OF THE MALE GAMETOPHYTE IN GINKGO BILOBA: THE SPERMATOGENOUS CELL

A light microscope and ultrastructural study was made of the pollen tube of Ginkgo biloba, with special emphasis given to the spermatogenous cell that gives rise to two motile sperms. Just prior to the mitotic division that results in the formation of two sperms, the spermatogenous cell consists of a large nucleus, two blepharoplasts, two large osmiophilic globules, and a conspicuous lipo-protein body. Other organelles in the cytoplasm include numerous electron-dense proplastids (with some lamellar development), mitochondria, small vacuoles, and lipid bodies. Ribosomes are present in abundance, but endoplasmic reticulum and dictyosomes are sparse. The nucleus, prior to mitosis, is relatively Feulgen-negative, due undoubtedly to the diffuse distribution of DNA. Each blepharoplast, the main organelle of interest, is nearly spherical, measures 3.5–4.5 μm in diam, and supports about 1,000 probasal bodies. The interior of a blepharoplast consists of an electron-dense matrix and of less dense regions which appear to be infiltrated by a network of microtubules. Each probasal body is composed of a cylinder of nine separate tubules (singlets) at the basal or proximal end. The cylinder becomes elaborated distally into nine pairs of subtubules (doublets) and then into nine sets of subtubules (triplets). A central tubule is present the entire length of the probasal body. Some of the subtubules, as well as microtubules from the interior of the blepharoplast, extend into the cytoplasm and probably constitute the “astral rays” as seen with the light microscope. Comparisons are made with other published accounts of the organization of blepharoplasts in plants and of centrosomes and centrioles in animals.     

蕨类植物分株紫萁精子发生过程中生毛体和多层结构的超微结构

应用电镜技术对蕨类植物分株紫萁(Osmunda cinnamomea L. var.asiatica Fernald)精子发育过程中的生毛体和多层结构的超微结构进行了研究.生毛体在幼精子细胞中出现,正在分化的生毛体略呈球状,球状体的中央由一团染色深的颗粒状物质构成,外围分化出若干柱状体.已分化的生毛体由柱状体分散或辐射状排列构成,呈球状,球体中心不含染色深的物质.多层结构位于精子细胞内的基体和巨大线粒体之间,刚形成时仅由片层构成,片层相互平行排列形成片层带.多层结构在分化中期由微管带、片层带和蚀斑三层构成.多层结构在分化末期又形成附属微管带、嗜锇冠和嗜锇层.微管带从多层结构长出,沿细胞核的表面伸展,并与核膜之间形成复合结构.基体由柱状体转变而成,它向两端生长,在远端产生鞭毛的轴丝,在近轴端形成楔状结构.本文首次详细阐明了原始薄囊蕨分株紫萁生毛体和多层结构发育的超微结构特点,并与其他蕨类进行了比较,发现其片层带出现在微管带形成之前.     

GIANT CENTRIOLE FORMATION IN SCIARA

Although somatic tissues of Sciara contain 9-membered centrioles, germ line tissues develop giant centrioles with 60–90 singlet tubules disposed in an oval array. Some 9-membered centrioles still may be seen in second instar spermatogonia. Each of these centrioles is associated with a larger "daughter" or secondary centriole at right angles to it. Most centrioles of second instar spermatogonia consist of 20–50 singlet tubules arranged in an oval, sometimes associated with an even larger secondary centriole. The more recently formed centriole of a pair is distinguishable from its partner by a concentric band of electron-opaque material inside its tubules. If a pair of centrioles at right angles to each other is pictured as a "T" formed by two cylinders, the secondary centriole is always the stem of the T; the primary centriole is the top. The two centrioles are oriented at the pole of the mitotic spindle so that the tubules of the primary centriole are parallel to the spindle axis. Each daughter cell receives a pair of centrioles and, during interphase, each of these centrioles gives rise to a new daughter centriole. A Golgi area of characteristic morphology is found in association with centrioles shortly after two new ones have formed. We conclude that in Sciara a centriole may give rise to a daughter morphologically different from itself. Whether the daughter is a 9-membered or giant centriole depends on the tissue type and stage of development.     

The fine structure of the spermatozoon of Pennaria tiarella (coelenterata)

Spermatozoa of the hydroid Pennaria tiarella were examined with the electron microscope. The anterior region is characterized by the presence of 30–40 membrane-bounded vesicles which lie anterior to the nucleus. These vesicles are apparently derived from the Golgi apparatus. The nucleus is conical in shape with a protrusion at the anterior end. Posteriorly it is indented by four radially arranged mitochondria. Lying within the fossa formed by the mitochondria are proximal and distal (filament forming) centrioles. The distal centriole is characterized by nine centriole satellite projections which emanate from its matrix. The tubules of the distal centriole are continuous with the alpha filaments of the tail. The tails are typical 9 + 2 flagella with 9 peripheral doublet (or alpha) filaments surrounding two central (or beta) filaments.     

Some ultrastructural aspects of spermatogenesis inLycopodium complanatum

Summary Analysis of thin sections shows that the blepharoplast in an early spermatid ofLycopodium consists of two basal bodies and the subtending spline apparatus. The latter has a 4-layered or Vierergruppe organization much like that reported for certain bryophytes and vascular plants. During the course of spermatid maturation the vertically lamellate S₂ stratum of the spline apparatus is transformed into a thinner layer of dense osmiophilic material, although the vertical lamellae of the S_3,4 strata persist. Rarely there occurred double Vierergruppen, presumably anomalous structures in which a single layer of spline microtubules is flanked on each side by two separate sets of S_2–4 strata. Longitudinal sections show that the basal bodies in an early spermatid lie parallel and close together with their microtubular triplets imbricating in the same direction. In late spermatids these basal bodies lie antiparallel and widely separated, their triplets now imbricating in opposite directions. This change is intrepreted to result from the relocation of one of the basal bodies to a position halfway around the cell by moving distal end first over the subtending spline. The basal bodies become invested by a globular-textured material not previously observed in plant spermatids. The role of this investing material is tentatively thought to be related to anchorage of the basal bodies.     

蕨类植物分株紫萁精子发生过程中生毛体和多层结构的超微结构

应用电镜技术对蕨类植物分株紫萁(Osmunda cinnamomea L. var．asiatica Fernald)精子发育过程中的生毛体和多层结构的超微结构进行了研究。牛毛体在幼精了细胞中出现,正在分化的生毛体略呈球状,球状体的中央由一团染色深的颗粒状物质构成,外围分化出若干柱状体。已分化的生毛体由柱状体分散或辐射状排列构成,呈球状,球体中心不含染色深的物质。多层结构位于精子细胞内的基体和巨大线粒体之间,刚形成时仅由片层构成,片层相互平行排列形成片层带。多层结构在分化中期由微管带、片层带和蚀斑三层构成。多层结构在分化末期又形成附属微管带、嗜锇冠和嗜锇层。微管带从多层结构长出,沿细胞核的表面伸展,并与核膜之间形成复合结构。基体由柱状体转变而成,它向两端生长,在远端产生鞭毛的轴丝,在近轴端形成楔状结构。本文首次详细阐明了原始薄囊蕨分株紫其生毛体和多层结构发育的超微结构特点,并与其他蕨类进行了比较,发现其片层带出现在微管带形成之前。     

The centrosome and bipolar spindle assembly: Does one have anything to do with the other?

In vertebrate somatic cells, the centrosome functions as the major microtubule-organizing center (MTOC), which splits and separates to form the poles of the mitotic spindle. However, the role of the centriole-containing centrosome in the formation of bipolar mitotic spindles continues to be controversial. Cells normally containing centrosomes are still able to build bipolar spindles after their centrioles have been removed or ablated. In naturally occurring cellular systems that lack centrioles, such as plant cells and many oocytes, bipolar spindles form in the complete absence of canonical centrosomes. These observations have led to the notion that centrosomes play no role during mitosis. However, recent work has re-examined spindle assembly in the absence of centrosomes, both in cells that naturally lack them and those that have had them experimentally removed. The results of these studies suggest that an appreciation of microtubule network organization, both before and after nuclear envelope breakdown (NEB), is the key to understanding the mechanisms that regulate spindle assembly and the generation of bipolarity.Key words: centrosome, centriole, mitosis, spindle, cell cycle, meiosis, plant cell, microsurgery     

CENTRIOLE REPLICATION : II. Sperm Formation in the Fern, Marsilea, and the Cycad, Zamia

Sperm formation was studied in the fern, Marsilea, and the cycad, Zamia, with particular emphasis on the centrioles. In Marsilea, the mature sperm possesses over 100 flagella, the basal bodies of which have the typical cylindrical structure of centrioles. Earlier observations by light microscopy suggested that these centrioles arise by fragmentation of a body known as the blepharoplast. In the youngest spermatids the blepharoplast is a hollow sphere approximately 0.8 µ in diameter. Its wall consists of closely packed immature centrioles, or procentrioles. The procentrioles are short cylinders which progressively lengthen during differentiation of the spermatid. At the same time they migrate to the surface of the cell, where each of them puts out a flagellum. A blepharoplast is found at each pole of the spindle during the last antheridial mitosis, and two blepharoplasts are found in the cytoplasm before this mitosis. Blepharoplasts are also found in the preceding cell generation, but their ultimate origin is obscure. Before the last mitosis the blepharoplasts are solid, consisting of a cluster of radially arranged tubules which bear some structural similarity to centrioles. In Zamia, similar stages are found during sperm formation, although here the number of flagella on each sperm is close to 20,000 and the blepharoplast measures about 10 µ in diameter. These observations are discussed in relation to theories of centriole replication.     

LEGOs® and legacies of centrioles and centrosomes

Centriole construction, now revealed by crystallography, proteomics, and imaging to be a sophisticated assembly of interlocking bricks, resembles LEGOs—albeit centrioles have remarkable dynamic capabilities, including self‐assembly and dis‐assembly, kinases and post‐translational modifications, self‐replication, and still mysterious mechanisms for transmission through each cell cycle and via the gametes during development. Centrioles are created by core proteins that aggregate to form unique ninefold‐symmetrical paracrystalline cylinders. The centrosome then coalesces as a cloud of pericentriolar material (PCM) around the centriole. Together they comprise the cell's microtubule organizing center (MTOC), which governs the shape, functions, and dynamics of the cell's microtubule (MT) arrays. This includes the meiotic and mitotic spindle apparatus for chromosome segregation, the accuracy of which is crucial for avoiding aneuploidies and resulting cancer, birth defects, or infertility. Centrioles’ replication and transmission mechanisms—and reduplication blocks—across cell cycles and generations, are only now becoming tractable to molecular analysis, which allows research to address questions about spindle assembly with neither centrioles nor centrosomes or de novo centriole formation. Here we discuss the latest insights into centriole and centrosome assembly and function and their transgenerational inheritance.     

THE STRUCTURE AND FUNCTION OF CENTRIOLES AND THEIR SATELLITES IN THE JELLYFISH PHIALIDIUM GREGARIUM

Testes of jellyfish Phialidium gregarium were fixed in 2 per cent OsO₄ in Veronal-acetate buffer at pH 7.4. Thin sections showed that in young spermatids the spindle fibers of the last maturation division are attached to satellites of the filament-forming centriole. In more mature spermatids this attachment is not observed. During the developmental phase, nine satellites can be observed emanating from the interspaces between the nine tubular triplets of this centriole. A circular region on each of the enlarged distal ends of the satellites attaches them to the cell membrane. The satellites apparently provide a firm anchor for the axial filament. Each of the epithelial cells covering the testis produces a single long flagellum. On the filament-forming centriole often a satellite can be observed to which tubules are attached. These tubules are 180 A in diameter and probably represent remnants of spindle fibers. It is suggested that the distal centriole has the ability to form several satellites or appendages at appropriate times during the cell cycle. These satellites are distinct from the daughter centrioles in that they are supportive structures: in certain phases of cell life, spindle fibers may attach to them, while in other instances the distal centriole and the flagellum it is forming are anchored by them.     

Identification of Microtubule-Organizing Centers in Interphase Melanophores of Xenopus Laevis Larvae In Vivo

The morphological characteristics of microtubule-organizing centers (MTOCs) in dermal interphase melanophores of Xenopus laevis larvae in vivo at 51-53 stages of development has been studied using immuno-stained semi-thick sections by fluorescent microscopy combined with computer image analysis. Computer image analysis of melanophores with aggregated and dispersed pigment granules, stained with the antibodies against the centrosome-specific component (CTR210) and tubulin, has revealed the presence of one main focus of microtubule convergence in the cell body, which coincides with the localization of the centrosome-specific antigen. An electron microscopy of those melanophores has shown that aggregation or dispersion of melanosomes is accompanied by changes in the morphological arrangement of the MTOC/centrosome. The centrosome in melanophores with dispersed pigment exhibits a conventional organization, and their melanosomes are situated in an immediate vicinity of the centrioles. In melanophores with aggregated pigment, MTOC is characterized by a three-zonal organization: the centrosome with centrioles, the centrosphere, and an outlying radial arrangement of microtubules and their associated inclusions. The centrosome in interphase melanophores is presumed to contain a pair of centrioles or numerous centrioles. Because of an inability of detecting additional MTOCs, it has been considered that an active MTOC in interphase melanophores of X. laevis is the centrosome. We assume that remaining intact microtubules in the cytoplasmic processes of mitotic melanophores (Rubina et al., 1999) derive either from the aster or the centrosome active at the interphase.     

Daughter Centriole Elongation Is Controlled by Proteolysis

The centrosome is the major microtubule-organizing center of most mammalian cells and consists of a pair of centrioles embedded in pericentriolar material. Before mitosis, the two centrioles duplicate and two new daughter centrioles form adjacent to each preexisting maternal centriole. After initiation of daughter centriole synthesis, the procentrioles elongate in a process that is poorly understood. Here, we show that inhibition of cellular proteolysis by Z-L₃VS or MG132 induces abnormal elongation of daughter centrioles to approximately 4 times their normal length. This activity of Z-L₃VS or MG132 was found to correlate with inhibition of intracellular protease-mediated substrate cleavage. Using a small interfering RNA screen, we identified a total of nine gene products that either attenuated (seven) or promoted (two) abnormal Z-L₃VS–induced daughter centriole elongation. Our hits included known regulators of centriole length, including CPAP and CP110, but, interestingly, several proteins involved in microtubule stability and anchoring as well as centrosome cohesion. This suggests that nonproteasomal functions, specifically inhibition of cellular proteases, may play an important and underappreciated role in the regulation of centriole elongation. They also highlight the complexity of daughter centriole length control and provide a framework for future studies to dissect the molecular details of this process.     

Breaking the ties that bind: new advances in centrosome biology

The centrosome, which consists of two centrioles and the surrounding pericentriolar material, is the primary microtubule-organizing center (MTOC) in animal cells. Like chromosomes, centrosomes duplicate once per cell cycle and defects that lead to abnormalities in the number of centrosomes result in genomic instability, a hallmark of most cancer cells. Increasing evidence suggests that the separation of the two centrioles (disengagement) is required for centrosome duplication. After centriole disengagement, a proteinaceous linker is established that still connects the two centrioles. In G2, this linker is resolved (centrosome separation), thereby allowing the centrosomes to separate and form the poles of the bipolar spindle. Recent work has identified new players that regulate these two processes and revealed unexpected mechanisms controlling the centrosome cycle.     

Spermatozoal Ultrastructure in Branchiostoma moretonensis Kelly, a Comparison with B. lanceolatum (Cephalochordata) and with other Deuterostomes

The spermatozoon of Branchiostoma moretonensis closely resembles that of B. lanceolatum and, though near the primitive sperm, allows recognition of a cephalochordate sperm type. This has: a bell-shaped acrosome; diffuse subacrosomal material not structured as an acrosome rod; sub-ovoidal nucleus with shallow anterior concavity, deep tubular posterior fossa (endonuclear canal) and condensed but lacunate chromatin; single asymmetrical, postnuclear mitochondrion almost completely or completely encircling the centrioles; mutually perpendicular proximal and distal centrioles of the triplet type, with the distal forming the basal body of the flagellum and the proximal (always?), as in B. moretonensis, with a spur-like extension (striated rootlet) into the nuclear fossa; flagellum tilted relative to the longitudinal axis (and endonuclear canal) of the nucleus; a 9 + 2 axoneme with hollow tubules and both dynein arms present on the doublets; and scattered glycogen granules, numerous around the distal centriole. The mitochondrion of B. moretonensis is C-shaped in transverse section, as in urochordates, but cephalochordate sperm resemble those of echinoderms, and specifically holothuroids, more closely. The occurrence of flagellar rootlets and composite mitochondria in various animal groups is discussed. The term paramorphy is proposed for parallel and convergent acquisition of an identical character: symparamorphy where acquisition is by parallelism and alloparamorphy where it is by convergence; the two terms represent, however, extremes of a continuum. Superficially similar but structurally different characters acquired by convergence are termed analogomorphis.     

Higher order structure of the PCM adjacent to the centriole

The centrosome is the major microtubule organizing center in most animal cells. This cytoplasmic organelle consists of two components : a mature centriole (or a pair of centrioles) and a mass of pericentriolar material (PCM). The PCM has been described as either a cloud of material that encases the entire centriole or as a cluster of proteins divided into two subsets, one that adheres to the lateral surface of the centriole and another that extends outward from this region as a cloud of material. In contrast to these protein distribution patterns, we demonstrated in a previous study that a subset of proteins present within the PCM is integrated together to form a tube (PCM tube) with an open and closed end that is duplicated in concert with centrosome duplication. The present study was undertaken to determine if this tubular conformation represents proteins that are confined to the surface of the centriole or if it represents a subset of proteins within the cloud of material that extends outward from the centriole. We document that : (1) the PCM tube represents a portion of the PCM directly associated with the centriole; (2) the PCM tube has a specific and reproducible relationship to the polar structure of the centriole; (3) the tube is a site of cytoplasmic microtubule organization, and has a structure that influences the initial pattern of microtubule assembly within the juxta-centriolar region; and (4) the PCM tube has a structural relationship with respect to the centriole, which allows the simultaneous expression of centriole- and PCM-based functions (e.g., ciliogenesis and cytoplasmic microtubule organization). Based on these findings, we propose a new model of the PCM at the centriole. This model highlights the role played by the proximal end of the centriole in the nucleation and organization of centriole-associated PCM, and indicates that the centrosome has an overall polarity in the region of the centriole.