MUCILAGE TUBE MORPHOLOGY OF THREE TUBE-DWELLING DIATOMS AND ITS DIAGNOSTIC VALUE1

Colonies of Berkeleya rutilans (Trent.) Grun., Navicula pseudocomoides Hendey and N. ramosissima (C. A. Ag.) Cleve were examined using light and electron microscopy to elucidate the structure of their mucilage tubes. Each species was found to have a characteristic tube construction which could be used for identification. A summary of features on which these species can be recognized was compiled.     

Identity and typification of Navicula hasta (Bacillariophyceae)

The name Navicula hasta is regularly applied to large, heavily silicified Navicula specimens with lanceolate outlines, protracted ends, and relatively coarse, radiate, and lineolate striae that are more irregularly spaced around the central area. As such, Navicula hasta has been reported from numerous localities such as brackish water fossil sediments, oligotrophic ancient lakes and postglacial lakes. When the data for distribution of Navicula hasta are summarized, this species appears to be geographically and temporally widespread with a broad range of ecologic tolerances. Published reports for size range show that diatomists during the last century have differing concepts of the species starting with the disagreement between the protologue and one of the figures given by Pantocsek. Furthermore, an informal ‘type’ of Navicula hasta reported in 1980 does not match the protologue and is not unambiguously based on material Pantocsek had in his hands when describing his new species. We studied published reports of Navicula hasta and investigated material reported to contain Navicula hasta and its allies from the type locality (Köpecz), one of the fossil deposits reported to contain N. hasta (Bodos) and from ancient and other younger lakes worldwide. From our analyses, we document the variation and underappreciated diversity within the Navicula hasta group and set the stage for formal recognition of new species in subsequent papers. Herein, we formally propose a lectotype from among the original illustrations and designate an epitype for Navicula hasta based on Pantocsek's Köpecz material. Additionally, two new species, Navicula krenneri spec. nov. and Navicula pseudohasta spec. nov. are described and a lectotype of Navicula superhasta Lange‐Bertalot and Metzeltin is proposed.     

Taxonomic studies on the diatom genus Navicula Bory. Navicula grevillii (CA.Ag.) Heiberg and N. comoides (Dillwyn) H. & M. Peragallo

Two closely related species of the diatom genus Navicula were investigated using light and electron microscopy. A hitherto unrecorded structure associated with the raphe of N. delognei Van Heurck is described.
Examination of type material revealed that the nomenclatural types of N. grevillii (CA.Ag.) Heiberg and N. comoides (Dillwyn) H. & M. Peragallo do not agree with the present understanding of these species, therefore alternative names were sought. All the synonyms of each species were traced and a table of synonymy is supplied. The names N. delognei and N. pseudocomoides are proposed as replacements for N. grevillii and N. comoides respectively and types are cited.     

THE ECOLOGICAL IMPLICATIONS OF GROWTH FORMS IN EPIBENTHIC DIATOMS 1

This paper evaluates the utilisation of space by epibenthic diatom cells, as a response to environmental variations. The aggregation pattern of five species of epibenthic diatoms was quantified and compared to provide evidence for the significance of cell motility as an adaptive mechanism for space occupation and monopoly. The epibenthic diatoms included (1) non-mobile colonial species forming either fan-shaped (Synedra tabulata (Ag)Kz.) or arborescent (Gomphonema kamtschaticum var. californicum Grun.) colonies; (2) slow-moving (Cocconeis costata Greg, and Amphora pusio Cl.), and (3) fast-moving (Navicula direct a (W. Sm.) Ra.) non-colonial species. The aggregation pattern of S. tabulata did not vary significantly among six different light intensities manipulated in nature. The major patterns of aggregation were identified using analysis of covariance and dummy-variable regression. Highly mobile N. directa are significantly less aggregated than the four other diatom species. Non-mobile and slow-moving species show a similar, highly aggregated pattern. The occurrence of two patterns of spatial dispersion indicates that growth forms bear far-ranging ecological implications with respect to colonization strategies, immigration, and possibly impact by grazers. An integrated model of growth form characteristics, biological properties, and ecological implications is presented for epibenthic diatoms.     

OBSERVATIONS ON NAVICULA ULVACEA,A RARE FOLIOSE MARINE DIATOM1

Navicula ulvacea (Berkeley ex Kütz.) P. T. Cleve, a marine colonial blade-forming diatom originally discovered in Scotland was collected at North Rustico, Prince Edward Island, during winter 1971. It formed dense stands of Porphyra-like fronds, ranging up to 60 mm long × 20 mm wide. These occurred under ice-covered water on rocks 1–2 m below zero tide. Comparisons made at the light and scanning electron microscope level compared favorably with original Aberdeen material. Clonal cultures established from single cells at first also formed blades, but rarely more than 10 mm long. In numerous serial subcultures made since 1971 no further blades have been formed. Instead, growth consisted of well-separated cells or short filaments scattered over the bottom and sides of the vessels. Older cultures developed prostrate sheets of filaments, embedded in a mucilagenous matrix and occasional bubble-like eruptions. Results of the cultures suggest that N. ulvacea probably occurs in a prostrate form in nature, perhaps more commonly than in the blade form. Literature records since the original collection in 1844 are brought together but are considered doubtful. A discussion on nomenclature is included.     

SEA ICE MICROBIAL COMMUNITIES. V. THE VERTICAL ZONATION OF DIATOMS IN AN ANTARCTIC FAST ICE COMMUNITY1

A distinct vertical zonation was observed among diatoms in a bottom congelation ice community at McMurdo Sound, Antarctica during the 1981 spring bloom. The bottom 20 cm of ice collected in December from four stations with variable snow cover was subdivided into 5 cm sections for analysis of algal distribution. Algal abundance was inversely related to the depth of snow cover, and generally decreased with increasing distance above the ice-water interface. Most diatoms, including the dominant species Nitzschia stellata Manguin, Amphiprora kufferathii Manguin and Fragilaria islandica var. adeliae Manguin showed peak abundance in the bottom 10 cm of the ice, where the proportion of living to empty cells was also highest. Two species, however, an Auricula Castracane sp. and Navicula glaciei van Heurck, reached highest concentrations at depths 10–20 cm above the ice-water interface. We considered two factors as contributing to the observed vertical zonation: (1) successive blooms at the ice-water interface become spatially stratified within the ice by further accretion below; (2) a differential growth of species occurs along physicochemical gradients within the ice column. A comparison of early versus late season profiles suggests the latter mechanism may prevail once ice accretion has ceased.     

LIGHT‐INDUCED MOTILE RESPONSES OF THE ESTUARINE BENTHIC DIATOMS NAVICULA PERMINUTA AND CYLINDROTHECA CLOSTERIUM (BACILLARIOPHYCEAE)1

Motility of estuarine epipelic (mud‐inhabiting) diatoms is an important adaptation to living in biofilms present within fine sediments. Motility allows cells to migrate within the photic zone in response to a wide range of environmental stimuli. The motile responses of two species of benthic diatoms to photon fluence rates and spectral quality were investigated. Cultures of Navicula perminuta (Grunow) in van Heurck and Cylindrotheca closterium (Ehrenb.) J. C. Lewin et Reimann both exhibited photoaccumulation at ～200 μmol · m^?2 · s^?1 and photodispersal from photon flux densities (PFDs) of ～15 μmol · m^?2 · s^?1. Photokinesis (changing cell speed) contributed toward photodispersal for both species, and red light (λ = 681–691 nm) was most effective at inducing this process. N. perminuta showed a phototactic (directional) response, with active movement in response to a light gradient. Although this response was exhibited in white light, these directional responses were only elicited by wavelengths from 430 to 510 nm. In contrast, C. closterium did not exhibit phototaxis under any light conditions used in this study. Motile benthic diatoms thus exhibit complex and sophisticated responses to light quantity and quality, involving combinations of photokinesis and phototaxis, which can contribute toward explaining the patterns of large‐scale cell movements observed in natural estuarine biofilms.     

MOLYBDENUM DEPENDENCE,NITRATE UPTAKE AND PHOTOSYNTHESIS OF FRESHWATER PLANKTON ALGAE1,2

The effects of molybdenum on photosynthesis and nitrate uptake by two species of freshwater algae, Navicula pelliculosa (Bréd.) Hilse and Chlamydomonas reinhardtii Dang. Were examined. Photosynthetic rates in cells of N. Pelliculosa deprived of Mo for 48 h were significantly lower than in nondeprived cells; the rates were 2.6 and 4.5 μg C/10⁶ cells/h, respectively. There was no significant reduction in photosynthetic rates of C. reinhardtii under the 2 treatments, the rates were both ca. 6.0 μg C/10⁶ cells/h. These observed rates in the 2 species were not altered by added Mo concentrations as high as 1.0 ppm. The chlorophyll a values in 48 h Mo-deprived cells were 1.15 μg/10⁶ cells compared to 1.57 μg/10⁶ cells provided with this metal. Molybdenum deficiency did not affect the chlorophyll a concentration in Chlamydomonas; the chlorophyll levels were 2.44μg/10⁶ cells. Nitrate uptake by Mo-deprived cells of N. Pelliculosa was significantly lower than in cells cultured in Mo; the rates were 0.087 and 0.238 μM NO_a⁻/10⁶ cells/h, respectively. Uptake rates by C. reinhardtii were similar with or deprived of Mo. The K_m values for No₃⁻ uptake were 14.9 and 148.0 μM for N. Pelliculosa and C. reinhardtii, respectively.     

EFFECTS OF METAL AND PETROLEUM HYDROCARBON CONTAMINATION ON BENTHIC DIATOM COMMUNITIES NEAR CASEY STATION,ANTARCTICA: AN EXPERIMENTAL APPROACH1

The effects of metals and petroleum hydrocarbons on benthic marine diatom communities were assessed using a manipulative field experiment at Casey Station, Antarctica. Uncontaminated, metal contaminated, and petroleum hydrocarbon contaminated sediments were deployed for 11 weeks during the 1999 austral summer. The treatments were deployed at three different locations: Brown Bay, which has elevated levels of anthropogenic contaminants, and two uncontaminated reference locations, O'Brien Bay and Sparkes Bay, the latter of which has naturally occurring high concentrations of some heavy metals. At each location, significant differences between the composition of diatom communities recruiting to control and petroleum hydrocarbon contaminated treatments were observed. Navicula directa (Smith) Ralfs occurred at lower abundances in the petroleum hydrocarbon contaminated treatments than in the control treatments. In contrast, Navicula cancellata Donkin occurred at higher relative abundances in both contaminated treatments relative to the control treatment. Interactions between treatment and location were also observed for several species, including Navicula glaciei Van Heurck. Significant differences in the overall community composition of diatom communities between control and metal contaminated treatments and metal contaminated and petroleum hydrocarbon contaminated treatments were only observed within Brown Bay. The location of deployment also had a significant influence on the composition of diatom communities. Brown Bay had higher abundances of Achnanthes brevipes Agardh but lower abundances of Navicula aff. cincta (Ehrenberg) Van Heurck than either reference locations. This experiment demonstrated that benthic diatom communities are sensitive to sediment contamination and would be suitable for future monitoring work within this and other areas of Antarctica.     

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.     

OBSERVATIONS ON NAVICULA THALLODES (BACILLARIOPHYCEAE), A BLADE-FORMING DIATOM FROM THE BERING SEA1

A thallus-forming diatom, Navicula thallodes Proschkina-Lavrenko, previously known only from the original collection at Bering Island (U.S.S.R.), has been found at Amchitka Island in the Aleutians, Alaska. The most remarkable observation of the present report is that N. thallodes may form blades up to 50 cm long, which to our knowledge is the greatest length reported for a colonial diatom. SEM observations of this diatom are presented for the first time.     

PHYSIOLOGICAL CHARACTERIZATION OF SIX LIPID-PRODUCING DIATOMS FROM THE SOUTHEASTERN UNITED STATES1

Six strains of diatoms from intertidal waters were isolated as part of the SERI Aquatic Species Program collection and screening effort: Amphiprora hyalina Greville, Cyclotella cryptica Reimann, Lewin & Guillard, Navicula acceptata Hustedt (two strains, NAVIC6 and NAVIC8), Navicula saprophila Lange-Bertalot & Bonik, and Nitzschia dissipata (Kütz.) Grunow. Among numerous algal strains isolated as part of this collection effort, these six strains showed rapid growth and elevated lipid content in preliminary screening experiments and were chosen for further physiological characterization. N. dissipata grew most rapidly at 25°C, whereas the other five strains grew best at 30–35°C. Salinity tolerance varied among strains, with maximal growth occurring at the following conductivities: 10–60 mS.cm^?1 (A. hyalina and N. acceptata NAVIC8), 10–35 mS.cm^?1 (C. cryptica), 20–45 mS.cm^?1 (N. acceptata NAVIC6), 10 mS.cm^?1 (N. saprophila), and 20–35 mS.cm^?1 (N. dissipata). The diatoms also differed in their utilization of nitrogen sources with A. hyalina growing optimally with either nitrate or urea; N. acceptata NAVIC6, with either nitrate or ammonium; C. cryptica, N. acceptata NAVIC8, and N. dissipata, with nitrate; and N. saprophila, with urea. Under optimal conditions, A. hyalina grew at 2.0 doublings. day^?1; C. cryptica grew at 3.0 doublings. day^?1. Each Navicula strain had a growth rate of 3.8 doublings. day^?1, and N. dissipata grew at 2.6 doublings.day^?1. All six strains had lipid contents in excess of 37% ashfree dry weight (AFDW) under nutrient-limited conditions, with N. saprophila having the highest lipid content at 48% AFDW.     

USING QUANTITATIVE PCR TO DETERMINE THE DISTRIBUTION OF A SEMICRYPTIC BENTHIC DIATOM,NAVICULA PHYLLEPTA (BACILLARIOPHYCEAE)1

Evidence has accumulated during the last decade showing that many established diatom morpho‐species actually consist of several semicryptic or truly cryptic species. As these species are difficult or even impossible to differentiate by microscopic analysis, there is virtually no information on how they behave in natural environments. In this study, we developed a quantitative real‐time PCR (qPCR) assay using TaqMan probes^® targeted to the internal transcribed spacer 1 (ITS1) to assess the spatial distribution and seasonal dynamics of an important component of the microphytobenthos of intertidal sediments. Navicula phyllepta Kützing is a brackish‐marine morpho‐species with a cosmopolitan distribution. Axenic clones of this species were isolated from natural assemblages of benthic diatoms at different intertidal stations in the Westerschelde estuary (The Netherlands). At least two distinct semicryptic species of N. phyllepta were present, as shown by differences in the quantity of DNA per cell, the ITS1 sequences and the copy number of ITS per cell. DNA and chl a concentrations extracted from sediment surface samples were closely correlated, showing that the DNA used for subsequent analysis mostly belonged to the microalgal community. The results of real‐time qPCR from sites throughout the estuary and over several seasons agreed well with microscopic counts. Additionally, the seasonal pattern of the two forms of N. phyllepta showed an overlapping, but unique distribution along the estuary.     

Examination of the type materials ofNavicula subtilissima Cleve (Bacillariophyceae)

Navicula subtilissima was described by Cleve as a new species from ‘a dry rivulet at the shore of Imandra, at the foot of Chibinä, Lapland’. However, the examination of the type slides and the unmounted original material using transmission and scanning electron microscopy revealed that there are at least three finely striated similar taxa distinguishable at the specific level. From these taxa, application of the nameN. subtilissima Cleve is established by selecting a lectotype; a second one is described as a new species,N. parasubtilissima, and the third is identified asN. subtilissima var.micropunctata Germain. The last one warrants a specific status by its fine structural differences in having continuous striae consisting of a single elongate areola occluded by a hymen with perforations arranging hexagonally. The occurrence of these three species is shown in Japanese collections.     

ICE‐BINDING PROTEINS FROM SEA ICE DIATOMS (BACILLARIOPHYCEAE)1

Sea ice diatoms thrive under conditions of low temperature and high salinity, and as a result are responsible for a significant fraction of polar photosynthesis. Their success may be owing in part to secretion of macromolecules that have previously been shown to interfere with the growth of ice and to have the ability to act as cryoprotectants. Here we show that one of these molecules, produced by the sea ice diatom Navicula glaciei Vanheurk, is a ～25 kDa ice‐binding protein (IBP). A cDNA obtained from another sea ice diatom, Fragilariopsis cylindrus Grunow, was found to encode a protein that closely matched the partially sequenced N. glaciei IBP, and enabled the amplification and sequencing of an N. glaciei IBP cDNA. Similar proteins are not present in the genome of the mesophilic diatom Thalassiosira pseudonana. Both proteins closely resemble antifreeze proteins from psychrophilic snow molds, and as a group represent a new class of IBPs that is distinct from other IBPs found in fish, insects and plants, and bacteria. The diatom IBPs also have striking similarities to three prokaryotic hypothetical proteins. Relatives of both snow molds and two of the prokaryotes have been found in sea ice, raising the possibility of a fungal or bacterial origin of diatom IBPs.     

OBSERVATIONS OF A TUBE-DWELLING DIATOM: NAVICULA HAMULIFERA (BACILLARIOPHYCEAE)1

Colonies of the tube-dwelling diatom Navicula hamulifera Grunow living on mangrove prop roots in Indian River. Florida and at La Parguera, Puerto Rico, were studied using light and electron microscopy. Observations of the tube morphology and cell structure of this diatom from fresh samples and cultures are described, as well as the ultrastructural morphology of its frustule. The formation of tubes by this diatom is reported for the first time. Comparisons are made with the closest species; Navicula delognei V.H. and Navicula pseudocomoides Hendey.     

新疆舟形藻属(硅藻门)中国新记录

报道了采自新疆的舟形藻属（硅藻门）中国新记录7个分类单位,如下所示：辐头舟形藻(Navicula capitatoradiata Germain)、辐指舟形藻(N. digitoradiata(Gregory) Ralfs)、瑞卡德提舟形藻(N. reichardtiana Lange-Bertalot)、针状舟形藻(N. spicula (Hickie) Cleve)、近小胎座舟形藻(N. subplacentula Hustedt)、琐细舟形藻(N. trivialis Lange Bertalot)和上凸舟形藻(N. upsaliensis(Grunow) Peragallo)。并对其分类学特征以及采集地生境进行了详细地描述。     

The production of extracellular carbohydrates by estuarine benthic diatoms: the effects of growth phase and light and dark treatment

Epipelic diatoms are important constituents of estuarine microphytobenthic biofilms. Field‐based investigations have shown that the production of carbohydrates by such taxa is ecologically important. However, limited information exists on the dynamics of carbohydrate production by individual species of epipelic diatoms. The production of low and high molecular weight extracellular carbohydrates in axenic cultures of five species of benthic estuarine diatoms, Cylindrotheca closterium (Ehrenberg), Navicula perminuta (Grun.) in Van Heurck, Nitzschia frustulum (Kütz.) Grunow, Nitzschia sigma (Kütz.) Grunow, and Surirella ovata (Kütz.) Grunow, were investigated. All species produced colloidal (water‐soluble) carbohydrates during growth, with maximal production occurring during stationary phase. During logarithmic growth, approximately 20% of extracellular carbohydrates consisted of polymeric material (extracellular polymeric substances [EPS]), but during stationary phase, EPS content increased to 34%–50%. Pyrolysis–mass spectrophotometry analysis showed differences in the composition of EPS produced during logarithmic and stationary phase. All species synthesized glucan as a storage carbohydrate, with maximum glucan accumulation during the transition from log to stationary phase. Short‐term labeling with ¹⁴C‐bicarbonate found that between 30 and 60% of photoassimilates were released as colloidal carbohydrate, with EPS consisting of approximately 16% of this colloidal fraction. When cells were placed in darkness, EPS production increased, and between 85 and 99% of extracellular carbohydrate produced was polymeric. Glucan reserves were utilized in dark conditions, with significant negative correlations between EPS and glucan for N. perminuta and S. ovata. Under dark conditions, cells continued to produce EPS for up to 3 days, although release of low molecular weight carbohydrates rapidly ceased when cells were dark treated. Three aspects of EPS production have been identified during this investigation: (1) production during rapid growth, which differs in composition from (2) EPS directly produced as a result of photosynthetic overflow during growth limiting conditions and (3) EPS produced for up to 3 days in the dark using intracellular storage reserves (glucans). The ecological implications of these patterns of production and utilization are discussed.