Population dynamics of an ice-associated diatom, Thalassiosira australis Peragallo, under fast ice near Syowa Station, East Antarctica, during austral summer

A clear shift from vegetative cells to auxospores and resting spores in Thalassiosira australis was observed in the water column and sinking fluxes under the fast ice near Syowa Station in the austral summer of 2005/2006. This is the first report of the auxosporulation by T. australis in situ. Resting spores were also observed in the sediment even before new spore formation, suggesting that T. australis can overwinter in the sediment. Heterotrophic dinoflagellates ingested and digested vegetative cells and auxospores but did not digest resting spores, suggesting a high tolerance of resting spores to grazing by heterotrophic dinoflagellates. We discuss the possible life history and overwintering strategy that T. australis uses in an Antarctic coastal area to cope with the unpredictable timing of sea ice growth and decay.     

Selection and viability after ingestion of vegetative cells, resting spores and resting cells of the marine diatom, Chaetoceros pseudocurvisetus, by two copepods

Feeding response of two copepods Neocalanus flemingeri and Calanus sinicus on cultures of three life-forms; vegetative cells, resting spores and resting cells, of Chaetoceros pseudocurvisetus was investigated. N. flemingeri fed heavily on the vegetative cells but scarcely responded to feed on the resting spores. C. sinicus showed significantly higher filtering rate on the vegetative cells and resting cells than on the resting spores. Survival of the three life-forms of C. pseudocurvisetus after gut passage of the copepods was also studied. The resting spores could germinate from fecal pellets of both N. flemingeri and C. sinicus; however, both the vegetative cells and the resting cells could not survive ingestion by the copepods. These results suggest that resting spore forming diatoms, such as C. pseudocurvisetus form spores which have a low nutritional value and during gut passage are largely indigestible due to the heavily silicified frustules and thus minimize the effects of grazing by copepods.     

EVIDENCE OF AN ORGANIC MATRIX FROM DIATOM BIOSILICA1

Most biominerals appear to be composites of organic material and mineral. Whether biosilica is such a composite is unresolved because of a lack of evidence for such organic components. We present evidence that organic material exists within diatom biosilica and can be extracted using HF/NH₄F solutions from frustules isolated from Cyclotella meneghiniana Kütz and diatomaceous earth. To eliminate organic casing on the silicified frustules as a source of organic materials, the casing was removed by oxidation of frustules with NaOCl before extraction. The removal of the casing was confirmed in that oxidized frustules no longer displayed the ability to be stained with ruthenium red and fluorescamine. Frustules examined with EDXA showed an emission peak from sulfur before treatment but no peak following treatment, indicating that oxidation removed organic sulfur. The organic material obtained from extracts of fresh frustules contained both soluble and insoluble components. Only soluble material was evident in extracts from diatomaceous earth. The soluble material appears to contain glycoproteins with relatively high levels of serine and glycine. The soluble proteins from fresh frustules also appear to be phosphorylated. Indirect evidence is presented that suggests the soluble proteins may contain regions of primary structure enriched in anionic amino acids. The soluble extracts differ from general cell contents when the two fractions are compared, suggesting that frustules contain specialized organic material. The identification of silica-specific organic material suggests that mineralization in diatoms may be in part matrix-mediated.     

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.     

Repeated evolution of uniparental reproduction in Sellaphora (Bacillariophyceae)

Diatoms possess a remarkable life cycle in which cell size decreases slowly during vegetative cell division and then increases rapidly via special expanding cells called ‘auxospores’, which are usually formed as a result of biparental sexual reproduction. However, auxospores are sometimes produced by single unpaired cells, i.e. uniparentally. We examined the nature of uniparental auxosporulation in Sellaphora and used a two-gene dataset to study phylogenetic relationships between uniparental and biparental Sellaphora demes and species; we tested whether uniparental reproduction has evolved once or repeatedly in the genus. In at least two of the uniparental demes auxosporulation occurred through autogamy (i.e. intra-tetrad mating within an undivided cell). Maximum likelihood phylogenies indicated four lineages of uniparental Sellaphora and significance tests of alternative topologies, in which combinations of uniparental Sellaphora were constrained to be monophyletic, coupled with likelihood reconstruction of ancestral character states, led to rejection of the hypothesis that uniparental auxosporulation evolved only once in the genus. Uniparentally reproducing lineages appear to arise not infrequently in diatoms but do not persist. Two small extranuclear bodies, apparently containing DNA and lying outside the chloroplast (one close to each pole of the cell), were revealed by DAPI staining.     

POLYMORPHISM IN MASTOGLOIA1

Cells in a -wild population of a species belonging to the diatom germs Mastogloia may form frustules of 2 strikingly different morphologic types during their vegetative life cycle. Total conversion from coarsely structured morphologic form to a more finely structured form takes place during a single division; hence no inter gradations between the 2 forms are evident. Intact frustules with 1 valve of each type leave no reasonable doubt of the ability to make the change.     

Sedimentation of biogenic siliceous particles in Antarctic waters from the Atlantic sector

Sediment traps were deployed in the Drake Passage, Bransfield Strait and west of South Orkney Islands (Powell Basin) during December 1980/ January 1981, December 1983, and between March and December 1983, respectively.Most of the trapped material is biogenic opal except in the lower half of the water column in Bransfield Strait where large amounts of resuspended aluminosilicates and quartz grains were present. Frustules and skeletons of siliceous microorganisms (diatoms, silicoflagellates, radiolarians, chrysophycean cysts), fragments and moults of crustaceans and tests of foraminifera were found. Quantitatively diatoms are the dominant constituent of the trapped biogenic material.Alteration of diatom assemblages in the water column is due to mechanical breakdown by grazing zooplankton. It mainly affects large frustules (e.g. Corethron criophilum Castracane) in the uppermost part of the water column. Dissolution of frustules occurs mostly at the sediment/water interface and leads to the enrichment of strongly silicified valves [e.g. Nitzschia kerguelensis (O'Meara) Hasle, Thalassiosira antarctica Comber (resting spores)].At the Bransfield Strait site a large part of biogenic opal was incorporated into fecal pellets of krill and copepods. The bulk of pellet content consists of fragmented diatom frustules (1–10 μm in size). Most intact valves found in the sediments have settled through the water column by means other than fecal pellet transport: e.g. settling as solitary particles or incorporated into or attached to “Marine Snow” or “Large Amorphous Aggregates”.     

Marine phytoplankton at the Weddell Sea ice edge: Seasonal changes at the specific level

Summary The Antarctic ice edge acts as a dynamic frontal system on the phytoplankton in the water column. Austral spring and autumn cruises to the Weddell Sea ice edge provided the opportunity to compare phytoplankton at the beginning of biological spring and at the end of biological autumn. The USCGC icebreakers Westwind (1983) and Glacier (1986) went into the sea ice, and the RV Melville (1983 and 1986) completed the transects in the adjacent open ocean. Field samples were observed alive on board ship to record different lifestages near the ice edge. In both seasons cell numbers were low under the ice, and single cells or short chains were the common growth habit. In spring in the open ocean, long chains of vegetative cells with large vacuoles and gelatinous colonies of diatoms and of prymnesiophytes dominated; in autumn in the open ocean close to the accreting ice edge, short chains, single cells, and resting spores were mostly packed with storage products. Enlarged cell diameters and auxospores also occurred near the ice cover in the autumn. Species from the following genera are included: the diatoms Leptocylindrus, Stellarima, Thalassiosira, Eucampia, Corethron, and Chaetoceros, the prymnesiophyte Phaeocystis, and the chrysophyte Distephanus.     

MOLECULAR EVIDENCE CONFIRMS SISTER RELATIONSHIP OF ARDISSONEA,CLIMACOSPHENIA, AND TOXARIUM WITHIN THE BIPOLAR CENTRIC DIATOMS (BACILLARIOPHYTA,MEDIOPHYCEAE), AND CLADISTIC ANALYSES CONFIRM THAT EXTREMELY ELONGATED SHAPE HAS ARISEN TWICE IN THE DIATOMS1

With the use of a new kit from Qiagen to amplify total genome quantity, DNA was bulked up from two diatoms that are difficult to grow (Ardissonea and Climacosphenia), and the nuclear SSU rRNA gene was successfully amplified. Results of Bayesian analyses showed that these diatoms are sister to Toxarium and belong to the bi‐ and multipolar centric diatoms. The results indicate that extremely elongate shape has arisen at least twice in diatoms, in the true pennates, and in the bipolar centrics. The two lateral pattern centers of Ardissonea and Climacosphenia likely represent a modified annulus that subtends ribs internally as well as externally. Studies of sexual reproduction are needed to determine whether Ardissonea, Climacosphenia, and Toxarium achieve their elongate shape by similar means to each other and to true pennates, that is, by controlling the expansion of the auxospores by sequential addition of silicified bands (to form a properizonium or perizonium).     

Morphology and ultrastructure of teratological forms of the diatoms Stephanodiscus niagarae and S. parvus (Bacillariophyceae) from Hamilton Harbour (Lake Ontario,Canada)

Teratological forms of Stephanodiscus niagarae Ehrenb. and S. parvus Stoermer & Håkansson were observed during a study of diatoms preserved in a radiometrically dated core from Hamilton Harbour (Lake Ontario, Canada). Morphological features and ultrastructures of both species were studied under the light and scanning electron microscope. The valve structure of abnormal forms of S. niagarae appears to be weakly silicified, especially in the central area. The shape of satellite pores are very irregular in comparison with the round shape of the normal specimen. The central fultoportulae are characterized as small tubes extending out of the external valve. Two types of abnormal frustules are present in the population of S. parvus studied. In one type valves are underdeveloped and only the siliceous layer and ribs are present, and in the other type the valves are strongly silicified and the areolae are almost completely occluded.The teratological forms of both species appeared in the core sediments after 1911, and both became the dominant components of diatom assemblages after 1970. Their occurrence and increased abundance coincides with records of heavy metal pollution in the harbour.     

NANOSTRUCTURE OF THE DIATOM FRUSTULE AS REVEALED BY ATOMIC FORCE AND SCANNING ELECTRON MICROSCOPY

The cell wall (frustule) of the freshwater diatom Pinnularia viridis (Nitzsch) Ehrenberg is composed of an assembly of highly silicified components and associated organic layers. We used atomic force microscopy (AFM) to investigate the nanostructure and relationship between the outermost surface organics and the siliceous frustule components of live diatoms under natural hydrated conditions. Contact mode AFM imaging revealed that the walls were coated in a thick mucilaginous material that was interrupted only in the vicinity of the raphe fissure. Analysis of this mucilage by force mode AFM demonstrated it to be a nonadhesive, soft, and compressible material. Application of greater force to the sample during repeated scanning enabled the mucilage to be swept from the hard underlying siliceous components and piled into columns on either side of the scan area by the scanning action of the tip. The mucilage columns remained intact for several hours without dissolving or settling back onto the cleaned valve surface, thereby revealing a cohesiveness that suggested a degree of cross-linking. The hard silicified surfaces of the diatom frustule appeared to be relatively smooth when living cells were imaged by AFM or when field-emission SEM was used to image chemically cleaned walls. AFM analysis of P. viridis frustules cleaved in cross-section revealed the nanostructure of the valve silica to be composed of a conglomerate of packed silica spheres that were 44.8 ± 0.7 nm in diameter. The silica spheres that comprised the girdle band biosilica were 40.3 ± 0.8 nm in diameter. Analysis of another heavily silicified diatom, Hantzschia amphioxys (Ehrenberg) Grunow, showed that the valve biosilica was composed of packed silica spheres that were 37.1 ± 1.4 nm and that silica particles from the girdle bands were 38.1 ± 0.5 nm. These results showed little variation in the size range of the silica particles within a particular frustule component (valve or girdle band), but there may be differences in particle size between these components within a diatom frustule and significant differences are found between species.     

Genotypic and phenotypic variation in diatom silicification under paleo‐oceanographic conditions

Diatoms have co‐evolved with the silicon cycle and are largely responsible for reducing surface concentrations of silicate in the ocean to their present levels. We quantify silicification in marine diatoms at a range of high silicate concentrations representative of environments found over their geological history. The species examined include Stephanopyxis turris, an ancient centric species found throughout the Cenozoic, Thalassiosira pseudonana and Thalassiosira weissflogii, two younger centric species, and two pennate ecotypes of Staurosirella pinnata isolated from different nutrient regimes. Frustule thickness and micromorphological structure are strongly affected by silicate concentration. All species become increasingly silicified with silicate concentrations at concentrations vastly in excess of surface ocean concentrations today. In contrast, the half‐saturation constant for silicate uptake for most modern diatoms is below 2 μm . Based on the results, we hypothesize that silicate uptake is multiphasic in diatoms and that multiple silicate transport systems may have evolved in response to decreases in surface silicate concentration over geological time. The oldest species examined is more heavily silicified than the more modern species, presumably reflecting the conditions under which it originated. Yet diversification in silicification can be rapid, as illustrated by greater silicification in onshore versus the offshore ecotype of the same modern species. This work suggests that silicification of fossil frustules may eventually provide a paleoproxy for surface silicate concentrations over the Cenozoic, although development of species‐specific calibrations will be necessary and the effects of a range of environmental conditions must be investigated.     

SEXUAL REPRODUCTION IN NAVICULA CRYPTOCEPHALA (BACILLARIOPHYCEAE)1

Homothallic sexual reproduction and auxosporulation were studied in monoclonal cultures and seminatural populations of the freshwater epipelic diatom Navicula cryptocephala Kütz. Gametangia paired via the girdle, one gamete was formed per gametangium (and hence one zygote per pair of gametangia), and gamete fusion took place without the formation of any copulation envelope or copulation canal. Superfluous nuclei from meiosis survived unusually long, so that gametes and young zygotes were probably functionally polyploid; later, all but two haploid nuclei degenerated. Expanded auxospores had a swollen center, but during formation of the initial valves, the auxospore contracted away from the perizonium to produce linear‐lanceolate valves. The pattern of reproductive behavior found in N. cryptocephala can be classified as type IIA2a auxosporulation in Geitler's system. The same type of zygote and auxospore formation seen in clonal cultures was observed in seminatural material from four lakes in Scotland and the Czech Republic. Variation in nuclear structure and auxosporulation in the N. cryptocephala species complex is discussed, as is the evolution of type II auxosporulation (one zygote per pair of gametangia) from type I auxosporulation (two zygotes per pair). The penalty of smaller numbers of zygote produced in type II may be outweighed by formation of larger auxospores (prolonging the vegetative phase) or more vigorous auxospores. The variation present among members of the N. cryptocephala complex indicates that biogeographical analyses based on use of the name N. cryptocephala, as performed recently to support the ubiquity hypothesis of protist distributions, are almost meaningless.     

Processing samples of benthic marine diatoms from Mediterranean oligotrophic areas

The processing of benthic diatoms is tedious and involves several potentially damaging steps for cells. Although the preservation of siliceous frustules is of paramount importance in the implementation of biotic indices, only few studies quantified treatment-induced cell losses. We assumed that commonly used treatments may lead to mechanical (centrifugation, sedimentation, boiling, sonication and mounting in Naphrax) and chemical (cold H₂O₂ digestion) damages on diatoms. We analysed the potential adverse effects of these treatments and the cleaning efficiency of H₂O₂ and incineration in order to find out the most suitable technique to process lightly silicified Mediterranean populations. Results showed that successive resuspensions of material after each concentration treatment (sedimentation and centrifugation) and low speed centrifugation did not alter the physical integrity of frustules. In contrast, boiling and sonication exhibited adverse effects especially on the preservation of large frustules and Naphrax mounting proved to be the most damaging step whatever the size of diatoms. For cleaning treatments, incineration provided the most satisfactory results and acted on a non-selective way as opposed to hydrogen peroxide which led to either a large number of non-cleaned frustules or dissolved valves. Our recommendations for processing samples of lightly silicified Mediterranean benthic diatoms include the use of low-speed centrifugations, dehydration at room temperature, incineration and dry mounting.     

Luminescence properties of a nanoporous freshwater diatom

Freshwater diatom frustules show special optical properties. In this paper we observed luminescence properties of the freshwater diatom Cyclotella meneghiniana. To confirm the morphological properties we present scanning electron microscopy (SEM) images. X‐ray diffraction (XRD) studies were carried out to visualize the structural properties of the frustules, confirming that silica present in diatom frustules crystallizes in an α‐quartz structure. Study of the optical properties of the silica frustules of diatoms using ultra‐violet‐visible (UV‐vis) spectroscopy and photoluminescence spectroscopy confirmed that the diatom C. meneghiniana shows luminescence in the blue region of the electromagnetic spectrum when irradiated with UV light. This property of diatoms can be exploited to obtain many applications in day‐to‐day life. Also, using time‐resolved photoluminescence spectroscopy (TRPL) it was confirmed that this species of diatom shows bi‐exponential decay. Copyright © 2011 John Wiley & Sons, Ltd.     

Studies on woloszynskioid dinoflagellates VI: description of Tovellia aveirensis sp. nov. (Dinophyceae), a new species of Tovelliaceae with spiny cysts

A new species of Tovellia, T. aveirensis, is described on the basis of light (LM) and scanning electron microscopy (SEM) of motile cells and resting cysts, complemented with transmission electron microscopy (TEM) of flagellate cells and phylogenetic analysis of partial sequences of the large subunit ribosomal rRNA gene. Both vegetative cells and several stages of a life cycle involving sexual reproduction and the production of resting cysts were examined in cultures established from a tank in the University of Aveiro campus. Vegetative cells were round and little compressed dorsoventrally; planozygotes were longer and had a proportionally larger epicone. Chloroplast lobes were shown by TEM to radiate from a central, branched pyrenoid, although this was difficult to ascertain in LM. The amphiesma of flagellate cells had mainly 5 or 6-sided vesicles with thin plates, arranged in 5–7 latitudinal series on the epicone, 3–5 on the hypocone. The cingulum had 2 rows of plates, the posterior row extending into the hypocone and crossed by a series of small projecting knobs along the lower edge of the cingulum. A line of narrow amphiesmal plates extended over the cell apex, from near the cingulum on the ventral side to the middle of the dorsal side of the epicone. Eight or 9 narrow amphiesmal plates lined each side of this apical line of plates (ALP). Resting cysts differed from any described before in having numerous long, tapering spines with branched tips distributed over most of the surface. Most mature cysts showed an equatorial constriction. Neither cysts nor motile cells were seen to accumulate red cytoplasmic bodies in any stage of the cultures. The phylogenetic analysis placed, with high statistical support, the new species within the genus Tovellia; it formed a clade, with moderate support, with T. sanguinea, a species notable for its reddening cells.     

Grazing impact and phenology of the freshwater sponge Ephydatia muelleri and the bryozoans Plumatella emarginata and Fredericella sultana under experimental warming

1. Both the clearance rates (CR) and abundances of the freshwater sponge Ephydatia muelleri and the bryozoans Plumatella emarginata and Fredericella sultana were investigated from autumn to spring under different temperature regimes. The experiments were performed in bypass channels of the River Rhine (Cologne, Germany) in which temperature could be manipulated. 2. The impact of temperature increase on CRs depends upon the grazer: E. muelleri showed a clear increase in CRs with increasing temperature whereas P. emarginata was not significantly affected by experimental warming. 3. Distinct differences in food preference were found for the sponge (which is an efficient grazer of bacteria and small algae) and for the bryozoan P. emarginata (which feeds primarily on large algae, and with no significant grazing on bacteria). 4. In contrast to their temperature‐related patterns in CR, respiration of both P. emarginata and E. muelleri increased with temperature between 19 and 32 °C, suggesting that the risk of experiencing energy deficiency at high temperatures due to increased metabolic rates is particularly high for the bryozoan. 5. A temperature elevation of 3 °C above the natural Rhine temperature resulted in a delay in the disappearance of active tissue and formation of resting stages for E. muelleri in autumn. This delay ranged from 8 (beginning of gemmulation) to 22 days (termination of gemmulation). In contrast, there was no distinct effect of warming on the disappearance of active zooids of the two bryozoan species in autumn. However, warming can positively affect the maintenance of active zooids during winter in F. sultana. In spring, the appearance of active zooids of P. emarginata was clearly stimulated by temperature elevations, whereas the hatching of both F. sultana and E. muelleri was hardly affected by warming. 6. The study demonstrated different patterns in the thermal ecology of the two freshwater bryozoans and the sponge in comparison to other filter feeders, particularly mussels. Such patterns need to be considered when predicting the impact of temperature on pelagic‐benthic coupling in aquatic habitats.     

HETEROSIGMA AKASHIWO (RAPHIDOPHYCEAE) RESTING CELL FORMATION IN BATCH CULTURE: STRAIN IDENTITY VERSUS PHYSIOLOGICAL RESPONSE 1,2

In this study, we examined the impact of environmental perturbation on the movement of the toxic bloom‐forming alga Heterosigma akashiwo (Hada) Hada ex Y. Hara et Chihara [syn. H. carterae (Hulburt) F.J.R. Taylor] between vegetative and resting cell phases of the life history. Resting state induction, in batch culture, was most effective when vegetative cells were subjected to low temperature (10° C) and darkness for extended time periods. Heterosigma cells in stasis had neither a cell wall nor scales but were surrounded by a calyx, most probably of polysaccharide composition. The resting cell was completely immobile, although both flagella remained attached. Heterosigma resting cells did not require a maturation period before successful activation to the vegetative state could occur. Cell division and motility were impacted sequentially during both the induction and activation phases of resting cell development. Our data show that Heterosigma had an obligate light requirement for resting cell activation. In replete medium, very low light fluences of 5 μmol photons·m ^{? 2}·s ^{? 1} were as effective as 60 μmol photons·m ^{? 2}·s ^{? 1} in the initiation of activation. Such sensitivity to extremely low light might give Heterosigma a competitive advantage for bloom formation in nature. Reduced nitrate levels significantly shortened the temporal transition of vegetative cells into the resting cell phase of the life history. Additionally, when resting cells induced in nitrate‐limited medium were activated under nitrate‐replete condition, the efficiency of the activation response was directly correlated to light availability. Both vegetative and resting cells maintained a haploid DNA complement. Rapid amplified polymorphic DNA (RAPD) analysis demonstrated variation in genetic identity among axenic Heterosigma strains. Strain identity influenced success in resting cell induction and survival in stasis. To date, no defined sexual cycle has been described. These observations are discussed in terms of population fitness. The data presented in this report provide a model algal system wherein the molecular events that govern long‐term stasis in an obligately autotrophic organism can now be assessed.     

Evolutionary significance of a middle Cambrian (Series 3) in situ occurrence of the pedunculate rhynchonelliform brachiopod Nisusia sulcata

Exceptionally preserved, silicified and articulated complete shells of the rhynchonelliform kutorginate brachiopod Nisusia sulcata are redescribed from the middle Cambrian (Series 3) Marjum Limestone, Utah. Cylindroid sausage‐like protrusions, emerging posteriorly between the valves, were originally interpreted as faecal in origin, but restudy under the SEM shows that these features represent silicified pedicles as they are attached in situ to other Nisusia. The Nisusia host most likely was alive at the time of attachment. Restudy of the pedicles of Nisusia provides new phylogenetic information on the anatomy of the earliest rhynchonelliforms. The silicified pedicles differ considerably from the pedicles of living crown group rhynchonelliforms in being strongly annulated, distally tapering, and were likely to have been rather more flexible. The Nisusia pedicles are more similar to the exceptionally preserved pedicles from other Cambrian rhynchonelliform brachiopods, including Kutorgina, Longtancunella and Alisina, but these emerge from the ventral apical foramen rather than from between the valves as in Nisusia. Although generally similar, these two types of pedicles are unlikely to represent homologous structures as Nisusia is provided with both an apical foramen (possibly larval attachment) and a posterior adult pedicle. The similarities may be explained by similar type of accretionary growth from two different types of epithelia. The Nisusia‐like pedicle appeared early within the kutorginates and rhynchonellates. The discovery of hollow spines in Nisusia sulcata further supports the generic assignation of the species.     

SEXUAL REPRODUCTION,MATING SYSTEM,AND PROTOPLAST DYNAMICS OF SEMINAVIS (BACILLARIOPHYCEAE)1

Cell division, the mating system, and auxosporulation were studied in the marine epipelic diatom Seminavis cf. robusta Danielidis & D. G. Mann. The interphase protoplast contains two girdle‐appressed chloroplasts, each with an elongate bar‐like pyrenoid, and also a central nucleus, located in a bridge between two vacuoles. Before cell division, the chloroplasts divide transversely and translocate onto the valves. The nucleus relocates to the ventral side for mitosis. After cytokinesis and valve formation, the chloroplasts move back to the girdle, showing a constant clockwise movement relative to the epitheca of the daughter cell. Seminavis cf. robusta is dioecious, and sexual reproduction is possible once cells are less than 50 μm. In crosses of compatible clones, gametangia pair laterally, without the formation of a copulation envelope, and produce two gametes apiece. The intensity of sexualization increases as cells reduce further in size below the 50‐μm threshold. At plasmogamy, the gametangia dehisce fully and the gametes, which were morphologically and behaviorally isogamous, fuse in the space between the gametangial thecae. The auxospore forms a transverse and longitudinal perizonium. After expansion is complete, there is an unequal contraction of the protoplast within the perizonium, creating the asymmetrical shape of the vegetative cell. Apart from this last feature, almost all characteristics exhibited by the live cell and auxospores of Seminavis agree with what is found in Navicula sensu stricto, supporting the classification of both in the Naviculaceae. Haploid parthenogenesis and polyploid auxospores were found, lending support to the view that change in ploidy may be a significant mechanism in diatom evolution.