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1.
Summary Cells of the centric diatomDitylum brightwellii were filmed undergoing cell division and valve secretion, and were fixed for transmission electron microscopy. Attention was directed particularly at the origin of the Labiate Process Apparatus (LPA).As reported previously (li andVolcani 1985 a), the nucleus, centrally situated during interphase, moves laterally to undergo mitosis against the girdle bands. We describe the spindle which splits up into numerous fibres of overlapped polar microtubules (MTs) by metaphase. The chromosomes are diffuse and the spindle elongates rapidly during anaphase. A complex of organelles is found at the poles and ill-defined, dense material extends to the nearby plasmalemma from prophase on. The two Silica Deposition Vesicles (SDVs) are initiated during anaphase close to the poles and by midcleavage, the dense LPA arises on each SDV close to dense polar material. After cleavage, the daughter protoplasts round up and the SDV, already containing a nascent valve, expands over the cleavage furrow. The labiate process, a long straight hollow tube of silica, is rapidly (ca. 25 minutes) secreted from directly under the LPA; a fibrous plug (polysaccharide?) always appears in the SDV immediately adjacent to the LPA during the initiation of this secretion. The ill-defined Microtubule-Organizing Center (MC) from the spindle pole remains close to the LPA and in it can be seen the tiny presumptive primordial spindle on the nuclear envelope.The raphe and the labiate process (LP), both highly differentiated apertures in the valve, probably function in a specialized form of the mucilage secretion involved in generation of movement in raphid diatoms, and in a simple form of movement in some centrics. Morphogenesis of the LP is associated with the LPA while differentiation of the raphe is almost associated with the MC; both MC and LPA have an intimate ontological relationship with the spindle pole and the postmitotic cytoskeletal system of MTs. This association also is seen in the formation of the LP in an araphid pennate,Diatoma (work in progress). Therefore, from functional, morphogenetic and ontogenetic observations, we support the proposal that the raphe of pennate diatoms arose from the LP of centric diatoms.  相似文献   

2.
Mitosis and valve morphogenesis in the pennate diatom Achnanthes coarctata (Bréb. in W. Sm.) Grun. are described. After cytokinesis, both daughter nuclei and their microtubule centers (MCs) are found near one side of the cell. Each new tubular silica deposition vesicle (SDV) arises centrally, forming a single rib running the length of the cell. Each MC then migrates around its nucleus and positions itself directly adjacent to the new SDV. The enlarging silicalemmas with their associated MCs, nuclei, microtubules (MTs) and microfilaments (MFs) appear in mirror image in the daughter cells. Both SDVs soon generate a second longitudinal rib alongside the first; the gap between the ribs ultimately becomes the future raphe fissure. The MC, MTs and nucleus are associated with each fissure. However, the subsequent behavior of the valve secreting machinery now becomes quite different in the daughter cells. In the cell that will form a raphid valve, the silicalemma, flanked by MFs, expands laterally in both directions over the cleavage furrow. Within the expanding SDV, silica secretion continues, eventually generating the structure of the mature valve, and during this phase the raphe fissure becomes delineated as in other raphid diatoms. In the other daughter cell, however, the MC and its MTs withdraw from the silicalemma, and the SDV moves laterally across the cleavage furrow until the double rib is at the corner of the cell. As silica is secreted into this expanding SDV, the raphe fissure completely fills in. This valve, therefore, lacks a raphe when mature and has a symmetry quite different from that of the valve formed in the other daughter cell. These events are compared with the course of morphogenesis described for other raphid diatoms.  相似文献   

3.
Valve morphogenesis in two Surirellae (S. ovalis Brebisson and S. robusta Ehrenberg) is described. Mitosis takes place at the broad end of the cell. After cleavage, a new Microtubule Center (MC) arises near each spindle pole and moves to the adjacent plasmalemma. Soon, a specific group of microtubules (MTs) extends from very near the MC around the periphery of the cell. Concurrently, the new tubular Silica Deposition Vesicle (SDV) grows around the periphery of the cell close to these MTs. A double rib of silica is rapidly formed inside the SDV; the space between the ribs becomes the raphe. Mitochondria line up along the MTs, and the SDV may be molded around these to create the canal raphe. Soon, the SDV expands in two directions to create the face and the mantle of the new valve. Meanwhile, each daughter nucleus, accompanied by the MC, moves to its interphase position at the center of the cell; this movement is colchicine-sensitive. As in several other pennate diatoms, an interruption in the raphe of the mature valve coincides with the initial position of the MC. The canal raphe thickens rapidly around the mitochondria; a rudimentary raphe fiber may be associated with the creation of a tiny curvature at the inner raphe fissure. As the SDV expands in the large S. robusta, the daughter cell protoplasts slowly shrink by plasmolysis, thereby creating the complex curved surface of the new valve surmounted by the arching canal raphes which are now quite rigid. In S. ovalis, the daughter cell protoplasts remain appressed and therefore the new valve surface is basically flat. The symmetry of Surirella is quite different from that of other pennate diatoms. However, the cytoplasmic events accompanying valve morphogenesis are similar in all important respects to those described in other raphid pennate diatoms, and clearly supports a naviculoid origin for this genus.  相似文献   

4.
Recent studies have led to a rapid increase in knowledge of auxospore formation in diatoms. However, these studies have been limited to centric and raphid pennate diatoms, and there is still very little information for the araphid pennate diatoms. Using LM and SEM, we studied the development of the auxospore and the initial cell of the marine epiphytic diatom Gephyria media Arnott. Auxospores were bipolar and curved in side view, as in many other pennate diatoms. SEM revealed many transverse perizonial bands, all of which were incomplete rings. There was an elongate, sprawling, silicified structure beneath the ventral suture of the transverse perizonial bands. This structure is presumably equivalent to the longitudinal perizonial band in other pennate diatoms, although we could not determine the homologous relationship between the two features. Scales were found both in the inner wall of the perizonium and around the primary perizonial bands. The presence or absence of scales may be of phylogenetic significance in diatoms, only during the final stages of auxospore formation because scales are found in early spherical stages. The distinctive finger‐like structures observed throughout all stage of G. media have not been observed before in the other diatom taxa.  相似文献   

5.
The deposition of siliceous valves during asexual reproduction of the pennate diatom, Navicula cuspidata Kütz., is described with emphasis on the cytoplasmic components involved. The events accompanying valve secretion are similar to those already known from other pennate species. After mitosis, the microtubule centre (MC) moves to the center of the cleavage furrow where silica deposition is initiated inside a tubular silicalemma, and it remains associated with the prospective central nodule during valve growth. Microtubules (MTs), emanating from the MC, run parallel to the prospective raphe and together with the raphe fibres, appear to be involved with raphe development. Multiple raphe fibres occupy the maturing raphe fissure, in contrast to the single fibre of Pinnularia viridis, P. maior and Hantzschia amphioxys. The fibers exhibit a periodic substructure and are often opposed to the silicalemma where they may inhibit silica deposition and control the shaping of the raphe fissure. In contrast with the above species, in N. cuspidata MTs are clustered strictly opposite the raphe and lose their association with the MC which degenerates before the valves are mature. The primary role of MTs may be the stabilization of the cytoplasmic region where initial silicification occurs. Mitochondria and endoplasmic reticulum are not involved in molding valve growth in this species. Evidence for vesicle involvement in silica transport and deposition was limited. The possible contributions provided by comparative studies on the ultrastructure of valve morphogenesis towards elucidating the control of valve formation and the taxonomy of diatoms are discussed briefly.  相似文献   

6.
After each division of a diatom cell, a new siliceous hypovalve is formed inside the silica deposition vesicle (SDV). We present the sequence of this early formation of the new valve in the pennate marine diatom Navicula salinarum (Grunow) Hustedt, visualized by using the fluorescent probe 2‐(4‐pyridyl)‐5‐((4‐(2‐dimethylaminoethylamino‐carbamoyl)methoxy)phenyl)oxazole (PDMPO). Our observations confirm that two‐dimensional expansion of the growing valve is a rapid process of no more than 15 min; three‐dimensional completion of the valve appears to be slower, lasting most of the time valve formation takes. The results are relevant to studies of the timing of molecular processes involved in valve formation (i.e. the bio‐ and morphogenesis of the SDV) in relation to uptake and transport of silicic acid. Use of this probe helps us to identify specific developmental stages for further detail analysis of diatom basilica formation, which eventually could lead to obtaining enriched SDV fractions.  相似文献   

7.
The pattern centre in valve morphogenesis is an annulus in centric diatoms or a sternum in pennate diatoms. The genus Rhaphoneis is currently placed within a lineage that diverges at the root of the pennate diatom clade in most molecular phylogenies, and its valves have a unique pattern to their striae, i.e. radiating from both apices, giving the impression that a pattern centre exists at both ends of the valve and virgae (ribs) formation proceeds centripetally. The present study, however, shows that the pattern centre is actually a linear sternum and the formation of virgae proceeds centrifugally, a pattern centre that is commonly found in most araphid diatoms. Thus, the hypothesis that valve morphogenesis based on a linear sternum and perpendicular virgae is a synapomorphy of pennate diatoms is supported. Our study also demonstrates that the pattern of valve formation can be observed by light microscopy with a direct mounting method when the specimen is relatively large, i.e. exceeding approximately 50 µm in valve length. An important advantage of the use of the direct mounting method is that it requires no repeated centrifugation steps for dehydration, steps necessary for observation by a scanning electron microscope, causing the loss and/or collapse of the specimen, particularly with fragile valves in the early stages of development.  相似文献   

8.
Valve morphogenesis in Proboscia alata Sundstrom was followed in living cells and during treatment with antiactin and antimicrotubule drugs. Once cleaved, sibling cells rounded up and retracted. Soon, a granular organizing center (OC) appeared adjacent to the stub of the initiated valve. Silicification started within a silicon deposition vesicle (SDV) adjacent to the OC. The elongating valve was initially tubular and sealed at one end, creating the proboscis of the conical valve. The edge of the SDV and thinnest region of the forming valve was lined by a sleeve of bundled microtubules (MTs) that terminated short of the older more rigid part of the valve. The growing proboscis of living cells treated with the anti‐MT drug oryzalin became grossly distorted. EM revealed dense material lining the growing edge of the SDV; immunofluorescence microscopy showed a ring of actin here. Applied to living cells, the antiactin drug cytochalasin D caused the very young proboscis to collapse; in older valves, the base of the proboscis expanded. Thus, valve morphogenesis appeared controlled by the MT cytoskeleton, keeping the proboscis straight while actin molded its conical outline. At the tip of the proboscis was a slit resembling a labiate process. Its morphogenesis involved striated fibers and two MTs, reminiscent of the fibers and MTs associated with raphe morphogenesis. In contrast to spine‐like processes that elongate by tip growth, the tip of the proboscis was formed first, and the consequent “antitip growth” suggests the tip was originally the center of the valve face.  相似文献   

9.
The diatom genus Eunotia is unusual among raphid diatoms in having a raphe system consisting of two short slits that are not integrated into the primary pattern center. This and other characteristics, particularly the presence of rimoportulae, are consistent with the hypothesis that Eunotia is a basal lineage within the raphid group. We studied auxosporulation in E. bilunaris (Ehrenberg) Mills and E. tropica Hustedt for comparison with other raphid pennate diatoms and with araphid pennates; E. bilunaris was studied in parental and F1 generations. Like araphid pennates, E. bilunaris and E. tropica are heterothallic. Clones of the same mating type did not interact sexually, and intraclonal sexual reproduction was absent or very rare. Clones retained the same sex throughout the life cycle, as shown by experiments using abrupt size reduction to produce clones of similar age but different size and using subclones derived from a single initial cell within six mitotic generations. Unlike in araphid pennate diatoms, in the Eunotia species the gametes are not visibly or behaviorally differentiated. Gametogenesis is merogenous, because the gametangium formed a supernumerary cell as well as a single gametic cell, both undergoing meiosis II to form a surviving functional nucleus and a nucleus that quickly degenerated. Plasmogamy is via papillae that grew out toward each other from the ends of the gametangia to create a copulation canal. After plasmogamy, the gametes moves bodily into the copulation canal, producing an elongate zygote, which expands to form a curved sausage‐like auxospore.  相似文献   

10.
Valve morphogenesis starts when the silica deposition vesicle (SDV) expands across a cleavage furrow covered by an unidentified layer, which may aid in its shaping. A labiate process (LP) is present only in the outer valve of terminal cells in the filament. Before these particular cells form setae, a layered "labiate process apparatus" (LPA) appears on the SDV in the exact center of the forming valve, near the microtubule center arising after cleavage. The LPA thereafter surmounts the lips of the LP as it forms. After the girdle bands separate slightly, two lateral protrusions develop in the corners of the cell. These nascent setae are lined internally by a cylindrical, fibrous band (sleeve), which assembles immediately ahead of the expanding edge of the SDV, very close to the plasmalemma. Then these protrusions, lined by the fibrous band, the SDV, and the forming silica wall, grow through two gaps in the girdle bands. The cytoplasm at the tip of the growing seta is naked. Immediately behind the tip, this fibrous band is adpressed to the plasmalemma and thereby apparently defines the diameter of the seta; it extends to internally ensheath the tipmost edge of the SDV for a short distance, like a tight-fitting inner sleeve. This structure is considered the major organelle involved in seta morphogenesis. Microtubules (MTs), while present, are variable in extent and disposition within the seta. Turgor pressure is considered irrelevant in driving seta growth. Instead, a new paradigm proposed for tip-growing cells generally, may apply to seta morphogenesis, as follows. If, as is suspected, the fibrous band contains actin, cycling of this actin (as in animal cells undergoing ruffling or filopodial extension) could drive seta extension via attachment of the band to the just-formed silica wall. The band is visualized as a molecular treadmill whose support base, the new wall, is being continually extended; extension is controlled and generated strictly at the tip.  相似文献   

11.
Diatoms have been classified historically as either centric or pennate based on a number of features, cell outline foremost among them. The consensus among nearly every estimate of the diatom phylogeny is that the traditional pennate diatoms (Pennales) constitute a well‐supported clade, whereas centric diatoms do not. The problem with the centric–pennate classification was highlighted by some recent analyses concerning the phylogenetic position of Toxarium, whereby it was concluded that this “centric” diatom independently evolved several pennate‐like characters including an elongate, pennate‐like cell outline. We performed several phylogenetic analyses to test the hypothesis that Toxarium evolved its elongate shape independently from Pennales. First, we reanalyzed the original data set used to infer the phylogenetic position of Toxarium and found that a more thorough heuristic search was necessary to find the optimal tree. Second, we aligned 181 diatom and eight outgroup SSU rDNA sequences to maximize the juxtapositioning of similar primary and secondary structure of the 18S rRNA molecule over a much broader sampling of diatoms. We then performed a number of phylogenetic analyses purposely based on disparate sets of assumptions and found that none of these analyses supported the conclusion that Toxarium acquired its pennate‐like outline independently from Pennales. Our results suggest that elongate outline is congruent with SSU rDNA data and may be synapomorphic for a larger, more inclusive clade than the traditional Pennales.  相似文献   

12.
Fucoxanthin–chlorophyll proteins (FCP) are the major light-harvesting proteins of diatom algae, a major contributor to marine carbon fixation. FCP complexes from representatives of centric (Cyclotella meneghiniana) and pennate (Phaeodactylum tricornutum) diatoms were prepared by sucrose gradient centrifugation and studied by means of electron microscopy followed by single particle analysis. The oligomeric FCP from a centric diatom were observed to take the form of unusual chain-like or circular shapes, a very unique supramolecular assembly for such antennas. The existence of the often disputed oligomeric form of FCP in pennate diatoms has been confirmed. Contrary to the centric diatom FCP, pennate diatom FCP oligomers are very similar to oligomeric antennas from related heterokont (Stramenopila) algae. Evolutionary aspects of the presence of novel light-harvesting protein arrangement in centric diatoms are discussed.  相似文献   

13.
The diatoms are one of the best characterised algal groups. Despite this, little is known of the evolution of the group from the earliest cell to the myriad of taxa known today. Relationships among taxa at the family or generic level have been recognised in some diatoms. However, relationships at higher taxonomic levels are poorly understood and have often been strongly influenced by the first appearances of key taxa in the fossil record. An independent assessment of relationships among the diatoms at these higher taxonomic levels has been made using rRNA sequence data to infer phylogenetic relationships. In this paper we present an analysis of 18S rRNA data from several chosen centric, araphid and raphid pennate taxa. The phylogenetic inferences from these 18S rRNA sequences are supported by evidence from the fossil record and evidence from ontogenetic data. Ribosomal RNA data indicate that both the centric and araphid pennate lineages may not be monophyletic.  相似文献   

14.
Prometaphase in two large species of diatoms is examined, using the following techniques: (a) time-lapse cinematography of chromosome movements in vivo; (b) electron microscopy of corresponding stages: (c) reconstruction of the microtubules (MTs) in the kinetochore fiber of chromosomes attached to the spindle. In vivo, the chromosomes independently commence oscillations back and forth to one pole. The kinetochore is usually at the leading edge of such chromosome movements; a variable time later both kinetochores undergo such oscillations but toward opposite poles and soon stretch poleward to establish stable bipolar attachment. Electron microscopy of early prometaphase shows that the kinetochores usually laterally associate with MTs that have one end attached to the spindle pole. At late prometaphase, most chromosomes are fully attached to the spindle, but the kinetochores on unattached chromosomes are bare of MTs. Reconstruction of the kinetochore fiber demonstrates that most of its MTs (96%) extend past the kinetochore and are thus apparently not nucleated there. At least one MT terminates at each kinetochore analyzed. Our interpretation is that the conventional view of kinetochore function cannot apply to diatoms. The kinetochore fiber in diatoms appears to be primarily composed of MTs from the poles, in contrast to the conventional view that many MTs of the kinetochore fiber are nucleated by the kinetochore. Similarly, chromosomes appear to initially orient their kinetochores to opposite poles by moving along MTs attached to the poles, instead of orientation effected by kinetochore MTs laterally associating with other MTs in the spindle. The function of the kinetochore in diatoms and other cell types is discussed.  相似文献   

15.
A marine araphid pennate diatom Plagiostriata goreensis is described from the sand grains of Goree Island, Dakar, Republic of Senegal, based on observations of fine structure of its frustule. The most striking feature of the species is its striation, which is angled at approximately 60° across the robust sternum. The other defining features of the species are its one highly reduced rimoportula and apical pores located at both ends of the valve margin. In the 18S rDNA phylogeny, the species appears as a member of a ‘small‐celled clade’ of araphid pennate diatoms that consist of Nanofrustulum, Opephora and Staurosira. The results of the phylogenetic analyses suggest that the distinct characters of the diatom; namely, oblique striae and apical pores, may have been acquired independently. However, it remains unclear whether the rimoportula of P. goreensis is a reduced state or P. goreensis acquired its morphologically curious rimoportula independently after the loss of an ancient rimoportula at the root of the small‐celled clade.  相似文献   

16.
Using a high-brilliance synchrotron X-ray source, combined small- and wide-angle X-ray scattering (SAXS and WAXS) was applied to study nanoscale characteristics, in particular pore size in the range of 3 to 65 nm, of a variety of unialgal cultures of centric and pennate diatoms, and of mixed diatom populations sampled in the field. Results of scattering analysis were compared with details of pore size, structure and orientation visible at the electron microscopic level. WAXS patterns did not reveal any crystalline phase or features of microcrystallinity (resolution 0.07 to 0.51 nm), which implies a totally amorphous character of the SiO2 matrix of the frustule material. SAXS data (resolution 3 to 65 nm) provided information on geometry, size, and distribution of pores in the silica. Overall, two pore regions were recognized that were common to the silica of all samples: the smallest (d less than 10 nm) regularly spaced and shaped spherically, the larger (up to 65 nm) being cylinders or slits. Apparently, at a nanoscale level diatomaceous silica is quite homologous among species, in agreement with the chemical principles of silica polymerization under the conditions of pH and precursor concentrations inside the silicon deposition vesicle. The final frustule "macro"-morphology is of course species-specific, being determined genetically. Synthetically-derived MCM-type silicas have a similarly organized pore distribution in an amorphous silica matrix as we found in all diatom species studied. We therefore suggest that organic molecules of a kind used as structure-directing agents to produce these artificial silicas play a role in the nucleation of the silica polymerization reaction and the shaping of pore morphology inside the silicon deposition vesicle of diatoms. Structure-directing molecules now await isolation from the SDV, followed by identification and characterisation by molecular techniques.  相似文献   

17.
The availability of extensive experimental data and remarkable intra- and interspecific variation in breeding behaviour make Achnanthes Bory sensu stricto an especially good model for studying the reproductive and population biology of pennate diatoms. In most Achnanthes species studied, auxospore formation is accompanied by biparental sexual reproduction, but we found uniparental auxosporulation in Achnanthes cf. subsessilis. Auxosporulation appears to be apomictic and follows contraction of the contents of unpaired cells and then a mitotic division, which is normally acytokinetic: one nucleus aborts before the cell develops into an auxospore. Rarely, both daughter nuclei survive and cytokinesis produces two auxospores (two auxospores per mother cell is highly unusual in pennate diatoms); one may abort. Expansion of auxospores is not accompanied by deposition of a transverse perizonium, but a longitudinal perizonium is produced and consists of a wide central strip (structurally similar to the araphid valve) and at least one narrow lateral strip. This newly discovered asexual lineage in Achnanthes is discussed in relation to other reproductive systems found in the genus, and also in relation to the ‘sex clock’ hypothesis concerning the adaptive significance of the diatom life cycle. Brief information on chloroplast division and nuclear dynamics over the cell cycle is also presented.  相似文献   

18.
A revision of the monoraphid pennate diatom genus Campyloneis Grunow was carried out based on LM and EM observations. The material examined originated from various herbarium collections and from extant epiphytic diatom communities on leaves of Posidonia spp. We also examined the generitype C. grevillei (Smith) Grunow and the fossil material of C. gheyselinchi Reinhold from which the author extracted the type. Our results clarified the fine structure of C. grevillei and C. gheyselinchi. Of the various varieties of C. grevillei, only the variety argus (Grunow) Cleve was retained. This differs from the nominate variety in the arrangement and shape of the areolae adjacent to the sternum of the araphid valve. The newly described taxon Campyloneis juliae De Stefano differs from all Campyloneis species in areolae ultrastructure and morphology of the valvocopulae. As for the fossil species C. gheyselinchi, the sternum valve areolae are similar to those of C. grevillei, but scarcity of frustules in the type material prohibited evaluation of its variability. For this reason we provisionally maintained its rank of species. The elaborate linking systems among the valvocopulae and valves in Campyloneis species appear to provide structural reinforcement against pressure from neighboring epiphytic diatoms and scouring of seagrass leaves.  相似文献   

19.
Oudot-Le Secq MP  Green BR 《Gene》2011,476(1-2):20-26
The mitochondrial genome of the raphid pennate diatom Phaeodactylum tricornutum has several novel features compared with the mitochondrial genomes of the centric diatom Thalassiosira pseudonana and the araphid pennate diatom Synedra acus. It is almost double the size (77,356 bp) due to a 35,454 bp sequence block consisting of an elaborate combination of direct repeats, making it the largest stramenopile (heterokont) mitochondrial genome known. In addition, the cox1 gene has a +1 translational frameshift involving Pro codons CCC and CCT, the first translational frameshift to be detected in an algal mitochondrial genome. The nad9 and rps14 genes are fused by the insertion of an in-frame sequence and cotranscribed. The nad11 gene is split into two parts corresponding to the FeS and molybdate-binding domains, but both parts are still on the mitochondrial genome, in contrast to the brown algae where the second domain appears to have been transferred to the nucleus. In contrast to P. tricornutum, the repeat region of T. pseudonana consists of a much smaller 4790 bp string of almost identical double-hairpin elements, evidence of slipped-strand mispairing and active gene conversion. The diatom mitochondrial genomes have undergone considerable gene rearrangement since the three lineages of diatoms diverged, but all three have kept their repeat regions segregated from their relatively compact coding regions.  相似文献   

20.
研究报道了采自中国四川九寨沟自然保护区长海的4种中心类硅藻种类, 其中新种1个: 长海小环藻(Cyclotella changhai sp. nov.), 中国新记录种1个: 克里特小环藻(C. cretica)和2种较少报道的种类: 可辨小环藻(C. distinguenda)和辐纹琳达藻(Lindavia radiosa)。通过光学显微镜和扫描电子显微镜对这4个种类形态学的特征进行观察, 并与相似种类进行比较。与长海小环藻相似的种类包括C. comensis Grunow, C. pseudocomensis Scheffler和C. costei Druart & Straub, 主要通过壳面中央区的不同形态进行区分-长海小环藻的中央区近平滑且具有不规则的凹陷。  相似文献   

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