STRUCTURE,LIFE HISTORY AND SYSTEMATICS OF RHOICOSPHENIA (BACILLARIOPHYTA). I. THE VEGETATIVE CELL OF RH. CURVATA1

Rhoicosphenia Grun. has been placed by some authors in the monoraphid group with Achnanthes Bory and Cocconeis Ehrenb., and by others near Gomphonema Ehrenb. In order to clarify the systematic position of the genus, the morphology and anatomy of the vegetative cells of Rh. curvata (Kütz.) Grun. were investigated using light and electron microscopy. The structure and formation of the two types of valve are described, and the heterovalvy shown to be of a different type from that of the monoraphids; on the basis of raphe, valve and girdle structure a close relationship between these and Rhoicosphenia is unlikely. Rhoicosphenia shows many resemblances to Gomphonema but the types of pore occlusion present, coupled with apparently slight differences in the mucilage-secreting structures and the girdle, suggest that classification in the same family is unwise. The cryptic asymmetry of the valves, and in particular of the raphe system, is noted and explained with reference to their formation; with respect to this asymmetry two configurations of the valves can occur (named cis and trans types) and the distribution of these in raphid genera is discussed briefly. In view of the lack of evidence in raphid diatoms supporting a classification of bands into copulae and pleurae, it is recommended that this practice be suspended.     

STRUCTURE,LIFE HISTORY AND SYSTEMATICS OF RHOICOSPHENIA (BACILLARIOPHYTA). II. AUXOSPORE FORMATION AND PERIZONIUM STRUCTURE OF RH. CURVATA1

Reproduction in Rhoicosphenia curvata (Kütz.) Grun. is isogamous. The two auxospores formed expand parallel to the apical axes of the gametangial cells. Expansion is bipolar and leads to the formation of a slightly curved, tapering cell, in which the initial valves are laid down. The perizonium consists of transverse and longitudinal bands. The transverse series, of 35 or so bands, is laid down centrifugally as the auxospore expands and can be classified into three groups on the basis of band morphology. All except the central band are open hoops, orientated so that their ends lie in the midline of the less convex, ventral side of the auxospore. The bands have fimbriate margins on one or both sides, and overlap one another from center to either pole. The longitudinal series includes 5 bands—a wide central band, with two on either side: again, the bands overlap one another from the center outwards. The initial epivalve of the new generation forms beneath the dorsal side of the auxospore, on the opposite side from the longitudinal perizonial series. Comparisons are made with other genera and the relevance of auxospore studies to an understanding of diatom morphogenesis is discussed.     

VALVE AND BAND MORPHOLOGY OF SOME FRESHWATER DIATOMS. III. PRE- AND POST-AUXOSPORE FRUSTULES AND THE INITIAL CELL OF MELOSIRA ROESEANA1

Frustules of a clonal culture of Melosira roeseana Rabenh. were examined with light and scanning electron microscopy. Vegetative valves in the post-auxospore (full size) stage exhibit a larger width/length ratio than those in the pre-auxospore (size-reduced) stage. Cells form chains by linking spines of adjacent valves which occur at the periphery of the valve face-mantle junction. Three or jour large pores occur at the center of the valve face, with the diameter of each pore tapering from the inner to the outer valve surface; these pores are often occluded by siliceous processes. Features of M. roeseana, not shown previously for Melosira, include a “stepped” mantle, on only one of the two valves resulting from the same cell division, flattened processes attached to short siliceous stalks on the valve face, disk-like processes on the mantle, and an open girdle band with up to eight antiligulae. Siliceous scales on the surface of the initial cell are remnants of the auxospore wall. The epivalve of the initial cell is larger in diameter than the hypovalve, and both valves lack linking spines and a step on the valve surface. The initial, cell epicingulum consists of only two bands; the hypocingulum has up to seven. Initial cells with four or more hypocingular bands divide to form new post-auxospore filaments. Melosira roeseana should not be included in the genus Melosira as it is presently defined by the type species, M. nurnmuloides C. Ag. Major differences include irregular linking spines, a closed pseudoloculate valve construction, and labiate processes on the valve face and mantle of M. nummuloides, compared with well-defined linking spines, a valve constructed of a basal siliceous layer perforated by poroid areolae, and labiate processes lacking on the valve of M. roeseana.     

STRUCTURE,LIFE HISTORY AND SYSTEMATICS OF RHOICOSPHENIA (BACILLARIOPHYTA). IV. CORRELATION OF SIZE REDUCTION WITH CHANGES IN VALVE MORPHOLOGY OF RH. GENUFLEXA1

Rhoicosphenia Grun. is a relatively isolated genus among the biraphid diatoms. Morphological changes in an isopolar member of the genus, Rh. genuflexa (Kütz.) Medlin, were investigated using light and scanning electron microscopy. The fully raphid valve showed changes in its flexure that could be correlated with size reduction during its life history from the initial cells to the smallest cells found in the population. Bands showed changes in number (from three to one) related to size reduction. Rh. genuflexa is morphologically similar to Rh. abbreviata (C. Ag.) Lange-Bert. (=Rh. curvata (Kütz.) Grun.), although the two are distinct taxa. These observations support previous contentions that Rhoicosphenia is a natural taxonomic grouping.     

SILICON DEPOSITION DURING THE CELL CYCLE OF THALASSIOSIRA WEISSFLOGII (BACILLARIOPHYCEAE) DETERMINED USING DUAL RHODAMINE 123 AND PROPIDIUM IODIDE STAINING1

The relatively non-toxic dye, rhodamine 123 (R123), was incorporated into the frustule of Thalassiosira weissflogii Grun. clone ACTIN in direct proportion to biogenic silica (BSi). R123 was used together with the DNA stain propidium iodide to track and quantify Si deposition during the cell cycle of T. weissflogii using flow cytometry. Silicon deposition was not continuous through the cell cycle. Deposition of the valves occurred during M phase. The hypocingulum was largely deposited during G1 with some suggestion of minor girdle band deposition during G2. Silicon deposition did not occur during S phase. Assuming that a complete frustule consists of an epivalve, epicingulum, hypocingulum, and hypovalve, then 40% of cellular BSi was contained within the cingulum of T. weissflogii with 60% present in the valves. These percentages correspond to 0.38 pmol Si in the two cingula and 0.57 pmol Si in the valves. Temporal differences in the timing of silicic acid uptake and deposition during the cell cycle of T. weissflogii suggested that deposition of both the new valves and the cingulum is supported by an internal pool of dissolved Si acquired during G2.     

Vallodiscus gen. nov., a new fossil resting spore morpho-genus related to the marine diatom genus Chaetoceros (Bacillariophyceae)

A new fossil marine diatom resting spore morphogenus, Vallodiscus Suto gen. nov., is described using samples from Deep Sea Drilling Project Site 338 in the Norwegian Sea, Sites 436 and 438 in the north‐west Pacific Ocean and the onland Newport Beach Section, California. Vallodiscus is characterized by a single ring of veins along the epivalve margin and a hypovalve covered with circular depressions of several sizes with gentle elevation. The morpho‐genus bears three new species and one new combination: Vallodiscus simplexus Suto sp. nov., Vallodiscus complexus Suto sp. nov., Vallodiscus lanceolatus Suto sp. nov. and Vallodiscus chinchae (Mereschkowsky) Suto comb. nov.     

POLYMORPHISM OF THE DIATOM PINNULARIA BREBISSONII IN CULTURE AND A FIELD COLLECTION1,2

Two clones of Pinnularia brebissonii (Kütz.) Rabh. var. brebissonii were established and maintained in logarithmic phase of growth. Initial length of the cells was 37 μm. As cell division occurred, the mean length of cells in each population decreased as predicted by the MacDonald-Pfitzer hypothesis; however, the decrease in mean length was not uniform throughout the growth period. This nonuniformity is probably caused by nonrandom division of cells in the population or by a changing increment of size reduction due to division. The initial increment of size reduction was calculated as 0.7 μm/division. The smallest, cells observed were 8 μm long. As cells decrease in length, cell volume decreases and the proportion of cells with aberrant valve structure increases. More than 90% of the valves were abnormal in a population with mean length of 14 μm. The abnormalities of structure involved the raphe, the central area and the striae.     

PERIZONIUM AND INITIAL VALVE FORMATION IN THE DIATOM NAVICULA CUSPIDATA (BACILLARIOPHYCEAE)1

This paper describes the perizonium and initial valve formation in Navicula cuspidata Kütz., based on light microscope (LM) and scanning electron microscope (SEM) observations. The perizonium consists of concentric over-lapping bands, laid down sequentially at the tips of the expanding biconical auxospore during its elongation. The central perizonial band has fimbriate edges and is considerably more rigid than the more distal bands. During auxospore elongation and the band secretion, the chloroplasts continuously oscillate between the two ends of the cell; this oscillation ceases once the elongation is complete. The initial valves, formed within the perizonium, are molded into the basically biconical shape of the perizonium except for a central flattening of each valve face. In contrast to the raphes in gametangial and vegetative valves which are surrounded by a smooth axial area, the raphes in initial valves lie within a raised ridge running along the apical axis of the valve. The regular pattern of apically oriented ridges on the outer surface of vegetative valves is also lacking on initial valves. Comparison of pore–pore spacing within striae of gametangial valves, initial values and post-initial valves (first division and vegetative cells) reveals that the pore–pore distance within striae is conserved at all sexual stages. However, the distance between striae is considerably larger in initial valves than in gametangial and post-initial valves. Vegetative interstriae spacing as well as the planar morphology of the valve face is regained at the first division of the initial cell. This suggests that the spacing between striae is dependent on the sexual stage of the cell during valve formation (i.e. not directly dependent on the cell size) and can be altered independently of the pore–pore spacing.     

VALVE MORPHOGENESIS AND THE MICROTUBULE CENTER IN THREE SPECIES OF THE DIATOM NITZSCHIA1

The relationship of cell organelles to valve morphogenesis was investigated in three species of Nitzschia. One, N. sigmoidea (Nitzsch) W. Sm., showed consistent ability to generate both nitzschioid and hantzschioid symmetry in daughter cells following cytokinesis; the other two maintained nitzschioid symmetry stably. From previous work with Hantzschia, a certain sequence of events could be anticipated in the cytoplasm. In two significant areas–the behavior of the Microtubule Center (MC) and its microtubule (MT) system, and the central origin of the silicalemma–not only were the results unexpected, but the three species showed fundamental differences among themselves. In N. sigmoidea, the silicalemma (and the future raphe region) arises centrally on the cleavage furrow, and after some lateral expansion, the silicalemmas and their associated organelles move in opposite directions in daughter cells, so that the raphe and the raphe canals end up along the girdle side of the cell as expected. However, the MCs never become associated with their silicalemma, remaining throughout near the girdle bands. In N. sigma (Kütz) W. Sm., the silicalemmas arise centrally and after lateral growth, move in opposite directions to generate nitzschioid symmetry. In this case, the MCs move to the vicinity of but never close to the silicalemmas, and follow them distantly during their lateral movement. In N. tryblionella Hantzsch, the new silicalemmas arise opposite one another, on one side of the daughter cells; each MC soon moves very close to its silicalemma, and remains thus through most of valve morphogenesis. Later, only one silicalemma/MC complex moves laterally, establishing the nitzschioid symmetry in both daughter cells. In all three species, as in Hantzschia, linear arrays of mitochondria aligned along MTs occupy the forming raphe canal, and microfilaments line the outer edge of the expanding silicalemma. The fibulae (the wall struts arching across the raphe canal) in Hantzschia always grow from the valve surface to the girdle surface of the forming valves. In these three Nitzschiae, this invariably happens in only one daughter cell of any pair; in the other, all the fibulae grow from the girdle surface to the valve surface. An explanation of these variations is proposed: that the morphogenetic machinery of Nitzschia and Hantzschia have a common origin, with present Nitzschiae having undergone considerable diversification at the intracellular level, causing the unstable cell symmetry exhibited by several modern species. Perhaps a taxonomic distinction between Hantzschia and Nitzschia lies in whether the morphogenetic machinery associated with valve morphogenesis moves laterally in the same or in opposite directions.     

Morphological investigations of the frustule, perizonium and initial valves of the freshwater diatom Achnanthes crenulata Grunow (Bacillariophyceae)

The present study clarifies the fine structure of the vegetative frustules, initial valves and perizonium of Achnanthes crenulata Grunow. The valves of the vegetative cell are distinctly linear‐lanceolate with an undulate margin. The valve face is quite flat and in girdle view is smoothly curved as in species of Gephyria (Bacillariophyceae). However, the valve face of the initial cells is slightly rounded and does not have an undulate margin. Furthermore, the rapheless sternum is centrally positioned along the apical axis of the araphid initial valve. As this taxon develops from auxospore to initial valve, it forms only longitudinal perizonial bands; no transverse bands arise. The perizonium consists of three silicified bands: one large, central longitudinal plate and two bands that underlie this plate; these two bands are either open or closed. This taxon has several conspicuous structures compared to other marine species of Achnanthes, but the structure of the perizonium supports the position of A. crenulata within Achnanthes sensu stricto.     

EFFECT OF SILICATE LIMITATION ON VALVE MORPHOLOGY IN THALASSIOSIRA AND COSCINODISCUS (BACILLARIOPHYCEAE)1

Coscinodiscus radiatus Ehrenb. and Thalassiosira eccentrica (Ehrenb.) Cleve were grown in a silicate-limited chemostat at silicate concentrations below 1 μg-atoms · l^?1. The resulting abnormal valves of C. radiatus lacked a thickened ring around the foramina; their pore membranes were thinner and their loculi shallower than those in normal cells. Abnormal valves of T. eccentrica had a fasciculate areolae pattern; they lacked a silica covering over the foramina and some tangential areolae walls. Neither abnormal valve could be termed a new species.     

VALVE MORPHOGENESIS IN THE PENNATE DIATOM NAVICULA CUSPIDATA1

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.     

Bewegungs- und Teilungsverhalten der Chromatophoren vonEunotia pectinalis var.polyplastidica und andererEunotia-Arten bei der Zellteilung

Summary The correlation between the start of chromatophore division and cell division is very different in various species ofEunotia. In some species the chromatophore division occurs before, in others after cell division. Eunotia pectinalis var.polyplastidica, with eight chromatophores per cell, represents an extreme type of behaviour in so far as two of the four plastids in each daughter cell prior to their division are shifted to the new hypovalve while the other two rest in situ. There occur two patterns of distribution of the four plastids, and that in the ratio 11, whereas the theoretically possible third pattern is never realized. The cause of that phenomenon is discussed. The division of the four plastids, in the meantime grown to full size, is performed not before they have reached their definitive equilibrium position at the epivalve or, respectively, at the hypovalve in twos. InEunotia pectinalis var.polyplastidica, by its mode of chromatophore division, the constant dissymmetric (right-left-handed) arrangement of the growing chromatophores, established inEunotia arcus, is not to be expected and is in fact not realized. InEunotia lunaris, however, the shift of the daughter chromatophores, in relation to the dorsiventrality of the cell, shows not only the same kind of dissymmetry as inEunotia arcus but also the same direction of shifting.The four and four chromatophores inEunotia pectinalis var.polyplastidica correspond to the two chromatophore plates in other species but the are not comparable with the numerous little chromatophore discs of Diatomaceae and other species the number of which decreases with the reduction of cell size.Cells ofEunotia pectinalis var.polyplastidica are capable to move by raphe action.

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Herrn Dr.Franz Berger zum 70. Geburtstag gewidmet.     

VALVE AND BAND MORPHOLOGY OF SOME FRESHWATER DIATOMS. IV. OUTER SURFACE MUCILAGE OF NAVICULA CONFERVACEA VAR. CONFERVACEA1

The development of the mucilage on the outer surface of Navicula confervacea (Kütz.) Grun., a raphed, filamentous diatom, was studied with scanning electron microscopy. This nonstructural cell wall material, present on the surface after critical-point drying and absent after acid cleaning, was of two types: strands and papillae. Strands were associated with the raphe system, areolae, elongated pores of the mantle, and all girdle sutures. Organic papillae were a common feature of valves, valvo-copulae and pleurae, but their origin and distribution could not be explained since they often occurred between the obvious openings in the frustule. Strands from the raphe and areolae may function in attaching terminal cells to a substrate and adjacent cells to each other. Other strands of the girdle arise from sutures during cell enlargement and continue to lengthen and intertwine until the individual frustules within a filament are obscured. Strands from sutures might originate from the advalvar row of pores of the girdle bands since these pores lie along the suture, but direct observation of this was not made. Secretion between, the bands also cannot be ruled out. Although mucilaginous papillae may sometimes occur at random on the entire surface of frustules, there is also a distinct, narrow multiseriate row of them around the edge of valves without marginal spines.     

Internal valves in populations of Meridion circulare from the A’er Mountain region of northeastern China: Implications for taxonomy and systematics

Abstract A population of Meridion circulare var. circulare (Greville) C.A. Agardh from Inner Mongolia was found to produce Innenschalen or internal spores. Examination of this population with light and scanning electron microscopy showed morphological differentiation between vegetative and spore morphologies. Vegetative valves typically bear costae and one rimoportula at the headpole. Spores lack costae and have two rimoportulae, one at the headpole and the other at the footpole. There is plasticity in the production of valve morphologies, and a variety of vegetative valve and spore combinations are evident. This population of M. circulare var. circulare has initial valves of over 90 μm in length, and all of the initial cells encountered are acostate and bear two rimoportulae. These observations suggest that either spores are the product of sexual reproduction, or that initial valves may be produced parthenogenetically in Meridion. Spores as products of the sexual process have not been reported in diatoms previously, and parthenogenesis in Meridion was reported previously but discounted in other published reports. The plasticity of valve morphologies expressed in M. circulare var. ciculare, between vegetative valves and spores (and back) across a short temporal period suggests that diatoms can alter their cell wall structure dramatically and quickly in response to external variables.     

VALVE AND BAND MORPHOLOGY OF SOME FRESHWATER DIATOMS I. FRAGILARIA CAPUCINA VAR. MESOLEPTA1,2

Acid cleaned cells from clonal cultures of Fragilaria capucina var. mesolepta Rabh. were examined with light and scanning electron microscopy. Recently isolated cells are linear-lanceolate in shape with a median constriction. After several transfers over 25 mo, cells exhibit size diminution resulting in small elliptically shaped valves. Adjacent valves are united to one another by interlocking marginal spines. Every valve has an apical pore field at each apex. A single labiate process is present infrequtently, appearing underdeveloped most often in size-reduced cells. The girdle region consists of two cingula, each composed of a series of underlapping bands. Each pleura in the series is a discontinuous ring with a central ligula. A survey of past ultrastructure studies on the freshwater Fragilariaceae reveals that the occurrence of the apical pore field and labiate process are likely key characteristics for the family. The apical pore field of Diatoma, Asterionella and Tabellaria is positioned on the valve face, whereas the apical pore field of F. capucina var. mesolepta is located on the valve mantle, the girdle region of F. capucina var. mesolepta is basically similar to that of Gomphonema parvulum (Kütz.) Grun.     

THE MORPHOLOGY AND INTEGRATION OF VALVES AND BANDS IN NAVICULA MUTICA VAR. MUTICA (BACILLARIOPHYCEAE)1

Navicula mutica (Kütz.) var. mutica was isolated from the air, cloned on agar, cultured in soil-water bottle, and studied with transmission and scanning electron micros-ropy. The frustules were lanceolate to ovoid with rounded apices, with the apical axis 8.5 ± 3.2 μ and the trans-apical and the transapical axis 3.6 ± 0.6 μm. Striae were composed of two or three puncta, and the mantle bore a single row of puncta aligned with the striae. The ends of the raphe turned away from an isolated punctual in the central area of the valve. The mantle puncta and one or two of the valve-face puncta in each stria opened into a series of transapical grooves in the interior of the valve, the grooves contributing to the appearance of striae in the light microscope. The interior of the mantle also possessed a pair of longitudinal grooves, discontinuous at the apices of the valves. An undulate advalvar margin of the valvocopula likely articulates along the interior longitudinal groove of the mantle. The projections of the undulate margin are perhaps positioned between the transapical grooves and along the longitudinal groove between the dentiform structures formed by the intersection of the double-grooved system. The girdle bands each had two (occasionally three) rows of pores. The pleurae margins were straight and not undulate.     

VEGETATIVE AND REPRODUCTIVE MORPHOLOGY OF EUCHEUMA ISIFORME (SOLIERIACEAE,GIGARTINALES, RHODOPHYTA)1

Eucheuma isiforme (C. Agardh) J. Agardh exhibits a combination of vegetative and reproductive features that distinguish it from other critically studied genera in the Solieriaceae. The development of the multiaxial thallus, emphasizing the arrangement of periaxial cells around each axial file; presence of reproductive nemathecia that contain carpogonial branches and auxiliary cells; and post-diploidization stages, including gonimoblast and pericarp initiation, stages of fusion cell formation, and carposporophyte development are described and illustrated for the first time in this species. The vegetative and reproductive features observed in E. isiforme are not diagnostic of any of the recently erected tribes in the Solieriaceae. Eucheuma appears most closely related to the Indian Ocean genus, Sarconema.     

GROWTH AND DIFFERENTIATION OF AXIAL AND LATERAL FILAMENTS IN BATRACHOSPERMUM SIRODOTII (RHODOPHYTA)1

Development of the vegetative gametophyte of Batrachospermum sirodotii Skuja was examined with light and both transmission and scanning electron microscopy. Patterns of wall growth were followed using the Calcofluor White ST pulse-chase method. Thallus structure was analysed in terms of the pattern of development of the apical, periaxial and pleuridial initials that generate the axial and whorled lateral filaments characteristic of Batrachospermum. Apical cells of axial filaments elongate initially by tip growth with the nucleus maintaining a distal position. Nuclear division is horizontal. One daughter nucleus migrates basipetally and a thin, convoluted annular septum and perforate-occluded pit connection are then formed. Elongating axial cells subsequently extend by wall deposition at the base of the cell. Periaxial cells are initiated laterally and elongate primarily by tip growth while the nucleus remains within the axial cell. The nucleus then migrates to the boundary between the initial and the axial cell, divides, and one daughter nucleus moves into the initial and the other back into the axial cell. A slightly irregular annular septum and simple-occluded pit connection are then formed. Pleuridial cell initials begin as terminal to subterminal protuberances on periaxial or pleuridial cells. They first extend by tip growth and later by bipolar band growth. The nucleus remains within the parent cell as the pleuridial initial expands and a narrow septal ring is formed between the two cells. It then migrates through the septal ring into the initial and divides transversely. One nucleus passes back into the parent cell and a thick, flat septum and perforate-occluded pit connection are formed. It is concluded that the potentially indeterminate axial filaments and the determinate lateral pleuridia represent distinct developmental types in Batrachospermum.     

DIATOM MINERALIZATION OF SILICIC ACID IV. KINETICS OF SOLUBLE Si POOL FORMATION IN EXPONENTIALLY GROWING AND SYNCHRONIZED NAVICULA PELLICULOSA1

Silicic acid taken up from the growth medium by Navicula pelliculosa (Bréb.) Hilse was shown to enter at least two compartments: i) soluble pools; ii) insoluble fraction comprised predominantly of the silica frustule. Soluble Si pools were extracted by a variety of agents from cells uniformly labeled for ten generations in medium containing ⁶⁸Ge-Si(OH)₄. 100 C water soluble and 0 C perchloric acid (PCA) soluble Si pools of 680 mM Si·l^?1 and 490 mM Si·l^?1 cell water represented 13 and 9%, respectively, of total, cell Si in exponential growth phase cells. Uniformly labeled cells synchronized by the combined synchronization technique accumulate at the cell cycle stage where silica frustule development is initiated. These cells contain water and PCA soluble pools of 10 nmol Si·10⁶ cells^?1 and-8.8 nmol Si·10⁶ cells^?1, respectively. On addition of Si(OH)₄, a rapid uptake ensues allowing the Si pool to expand 2.5-fold, apparently to provide precursors of the silica frustule.