THE EFFECT OF IRRADIANCE ON THE GROWTH OF DIFFERENT FORMS OF FRESHWATER DIATOMS: IMPLICATIONS FOR SUCCESSION IN ATTACHED DIATOM COMMUNITIES

The growth rates of 12 strains of attached freshwater diatoms were examined as a function of irradiance. The results were used to determine physiological parameters in the growth–irradiance relationships. Values of G _max (maximum growth rate) and I  _k (irradiance at half of G _max) correlated with successional phase. The development of attached diatom communities appears to proceed from high irradiance species to low irradiance species that appear later in the successional sequence. Thus, efficiency of light use is linked to successional strategy, as has been documented for terrestrial plants.     

THE ROLE OF DIATOMS IN STROMATOLITE GROWTH: TWO EXAMPLES FROM MODERN FRESHWATER SETTINGS1

Some modern laminated and calcified stromatolitic structures are partially or completely formed by eukaryotes. Diatom populations in freshwater environments with elevated ionic concentrations contribute to calcite precipitation, and the formation of distinctive mineral-rich stromatolitic laminae. Two types of stromatolite-forming diatom populations were observed. In the first example, in stromatolies growing on a quarry ledge near Laegerdorf, North Germany, calcite crystals with biogenic imprints form around polysaccharide stalks of the diatom Gomphonema olivaceum var. calcarea (Cleve) Cleve-Euler. These individually precipitated crystals eventunally become cemented together in layers, forming rigid, laminated stromatolitic deposits which drape over the quarry ledge. In the second example, in stromatolites forming in a shallow stream near Cuatro Ciénegas, Coahuila, Mexico, diatomaceous laminae also form by the accumulation of carbonate particles in a matrix of diatoms and their extracellular polysaccharide products. These laminae become thick enough to drape over individual stromatolite heads. The diatoms responsible for these deposits are Amphora aff. A. Katii Selva, Nitzschia denticula Grun., and six other species. At Cuatro Ciénegas, in addition to the diatomaceous laminae, carbonate-rich cyanobacterial layers, dominated by two cyanobacterial species with different fabrics and porosities, are also present and contribute substantially to the growth of the stromatolites. In both the Laegerdorf and Cuatro Ciénegas examples, entire stromatolites or thick laminations on stromatolites are built by a small number of diatom species which produce copious amounts of extracellular stalk, gel, and sheath material, a property they share with cyanobacterial stromatolite builders.     

THE ROLE OF DIATOMS IN STROMATOLITE GROWTH: TWO EXAMPLES FORM MODERN FRESHWATER SETTINGS1

Some modern laminated find calcified stromatolitic structures are partially or completely formed by eukaryotes. Diatom populations in freshwater environments with elevated ionic concentrations contribute to calcite precipitation, and the formation of distinctive mineral-rich stromatolitic laminae. Two types of stromatolite-forming diatom populations were observed. In the first example, in stromatolites growing on a quarry ledge near Laegerdorf, North Germany, calcite crystals with biogenic imprints form around polysaccharide stalks of the diatom Gomphonema olivaceum var. calcarea (Cleve) Cleve-Euler. These individually precipitated crystals eventually become cemented together in layers, forming rigid, laminated stromatolitic deposits which drape over the quarry ledge. In the second example, in stromatolites forming in a shallow stream near Cuatro Ciénegas, Coahuila, Mexico, diatomaceous laminae also form by the accumulation of carbonate particles in a matrix of diatoms and their extracellular polysaccharide products. These laminae become thick enough to drape over individual stromatolite heads. The diatoms responsible for these deposits are Amphora aff. A. katii Selva, Nitzschia denticula Grun., and six other species. At Cuatro Ciénegas, in addition to the diatomaceous laminae, carbonate-rich cyanobacterial layers, dominated by two cyanobacterial species with different fabrics and porosities, are also present and contribute substantially to the growth of the stromatolites. In both the Laegerdorf and Cuatro Ciénegas examples, entire stromatolites or thick laminations on stromatolites are built by a small number of diatom species which produce copious amounts of extracellular stalk, gel, and sheath material, a propertuy they share with cyanobacterial stromatolite builders.     

DISTRIBUTION OF DIATOMS IN THE PLANKTON OF YAQUINA ESTUARY,OREGON1

The distributional patterns of diatoms in the plankton of the Yaqnina Estuary, Oregon, were, investigated and related to selected climatic and hydrographic factors. Distribution was strongly influenced by seasonal patterns of rainfall resulting in the introduction of a large volume of fresh water into the estuary during fall and winter. Plankton assemblages in spring, summer and fall had fewer diatom species and exhibited a more rapid rate of change in species composition than in winter. Winter assemblages were further characterized by many pennate diatoms, apparently dislodged from the benthos during periods of high freshwater discharge and silt loads. A statistical measure of community difference indicated an increase in taxonomic homogeneity among assemblages throughout the estuary with the onset of the rainy season in late fall and a gradual transition to a more heterogeneous system again during late spring. Canonical correlation ordered 20 prominent diatom taxa along the salinity gradient and identified possible relationships among certain taxa and selected environmental variables, namely visible light energy and temperature. Redundancy in the species data given the environmental data was only 40%, emphasizing the difficulty in demonstrating a quantitive relationship between plankton dynamics in the field and concurrent measurements of chemical and physical variables.     

EXAMINATION OF TWO DIMENSIONAL SPATIAL PATTERN OF PERIPHYTIC DIATOMS USING AN ADHESIVE SURFICIAL PEEL TECHNIQUE1

A simple surficial peel technique using adhesive tape was developed for quantitative removal of haptobenthic diatom communities from topographically simple substrata. The method combines high removal efficiency with low peel distortion, permitting the use of spatial statistics to test whether populations are distributed in the peel randomly or form aggregated or uniform patterns. Using this technique, the microdistribution of Cocconeis placentula Ehr. on a smooth acrylic rod was examined. Using conventional nearest neighbor analyses, a clonal population of C. placentula. characterized by an indentation of the value margin, was significantly aggregated, whereas the overall C. placentula population was uniform or aggregated depending on whether the method of analysis allowed for cell size. Using refined nearest neighbor analysis, the indented population was aggregated, and the overall population was random at distances greater than cell size. The results suggest that the indented clone was weakly motile following cell division and that its directional bearing was random.     

COMMUNITY STRUCTURE DYNAMICS OF EPILITHIC AND EPIPHYTIC DIATOMS IN A MISSISSIPPI STREAM1

Epilithic and epiphytic diatoms were collected monthly from 18 June 1979 through 19 May 1980 from both shaded and unshaded sites of Sessums Creek, a shallow, oligotrophic and strongly alkaline stream in northeastern Mississippi. The dominant macroalgae were filamentous green algae belonging to the non-mucilage producing genera Cladophora and Rhizoclonium. Ninety taxa were identified with Navicula minima Grun. and Nitzschia dissipata (Kütz.) Grun. accounting for 50% of all valves counted. Other important taxa were Navicula menisculus var. upsaliensis (Grun.) Grun., N. cryptocephala var. veneta (Kütz.) Rabh., Achnanthes minutissima Kütz. and Cymbella turgida (Greg.) Cl. With few exceptions, the more dominant taxa were equally abundant in the shaded and unshaded sites and also in the epilithon and epiphyton. Species diversity (H') and the number of taxa in a sample in all four habitats showed an identical pattern, being lowest in winter and highest in fall. No one pair of habitats was more structurally similar than any-other pair when compared by a similarity index (SIMI). Apparently, differences in light intensity and substrate type had little effect on diatom distribution in Sessums Creek and it was concluded that the locality sampled supported a single, nearly homogeneous diatom community.     

PRODUCTION OF EPIBENTHIC DIATOMS IN A SOUTHERN CALLIPORNIA IMPOUNDED ESTUARY1, 2

In situ epibenthic diatom productio was determined for four sites within Bolsa Bay (Orange Coutry), California, to assess the effect of estuative impoundment by a tidal gate. Mean annual values of 26 and 41 mg C m^?2 h^?1 were found in the finner sediment of the outer bay whereas mean annual values of 41 and 59 mg C m^?2 h ^?1 were found in the coarser, more stable sediment of the inner bay. Low production in the vicinity of the mouth of a flood control channel is thought to be caused by heavy metal inhibition in conjuction with sediment instability, a shl/allow photic zone and heavy grazing pressure. Seasonal production peaks in spring and falll were found to be closely correlated with the concentration of nitrate and silicate but not with phosphate. This study shows that even though Bolsa Bay has been cut off from the sea, drained and subjected to fill, it is as productive as other coastal wetlands throughout the world.     

INFLUENCE OF CELL VOLUME ON THE GROWTH AND SIZE REDUCTION OF MARINE AND ESTUARINE DIATOMS1

The maximal growth rate (μ_max) of 19 marine and estuarine diatoms decreased with increasing cell volume (V). The relationship between log μ_max (Y) and log V (X) was calculated. Statistical analyses showed that the slope of the equation was not significantly different from those obtained by other researchers and that the 95% confidence intervals of mean μ_max at cell volumes of 10³–10⁵μm³ were not significantly different from those cited in most studies. A new regression line for diatoms was calculated as follows: log μ_max= 0.47–0.14 log V; r =–0.69. The rate of size reduction per generation of the 19 diatom species ranged from 0.03 to 0.87 μm per generation. The rate increased with increasing cell length and cell volume and with decreasing maximum division rate. Statistical analyses showed that the rate was closely related to the cell volume and to the reciprocal of the growth rate. The relationships between maximal growth rate and cell volume and between rate of size reduction and cell volume showed that a diatom with a large volume had a smaller maximal growth rate and a larger rate of size reduction than a diatom with a small volume. The estimates using the equation for the regression line between the rate of size reduction and the reciprocal of maximum division rate indicated that a diatom with a high maximum division rate would need more generation equivalents for a certain size reduction than a diatom with a low maximum division rate, but the periods required for reduction would be approximately equal irrespective of maximum division rate.     

EFFECT OF POLYCARBONATE CONTAINERS ON THE GROWTH OF TWO SPECIES OF MARINE DIATOMS1

Two species of marine diatoms [Skeletonema costatum (Greville) Cleve and Thalassiosira pseudonana (Hustedt) Hasle and Heimdal] were grown in glass and polyarbonate containers. S. costatum exhibited a signzJicantly lower exponential growth rate and maximal yield and a signajcantly longer lag phase when grown in polycarbonate. Exponential growth rate and maximal yield of T. pseudonana was significantly reduced (P < 0.05 in all cases). This study suggests that a difference in diatom growth between glass and polyarbonate containers might arise in certain cases. However, such a difference may not be detectable with all biomass measurement techniques or with low within-treatment replication.     

DIATOMS (BACILLARIOPHYTA) OF ISOLATED ISLANDS: NEW TAXA IN THE GENUS NAVICULA SENSU STRICTO FROM THE GALáPAGOS ISLANDS1

The diatoms (Bacillariophyta) from a coastal lagoon from the Diablas wetlands (Isla Isabela, the Galápagos Islands) were studied in material from surface samples and a sediment core spanning the past 2,700 years in order to examine evidence of diatom evolution under geographic isolation. The total number of taxa found was ～100. Ultrastructural variation in valve morphology between members of Galápagos taxa was used to describe 10 species from the genus Navicula sensu stricto, which are new to science. Four taxa: N. isabelensis, N. isabelensoides, N. isabelensiformis, and N. isabelensiminor, shared several key characteristics that may be indicative of a common evolutionary heritage; these species therefore provide possible evidence for the in situ evolution of diatoms in the Galápagos coastal lagoons. Shared morphological characteristics include: (i) stria patterning in the central area, (ii) an elevated and thickened external raphe‐sternum, (iii) external central raphe endings that are slightly deflected toward the valve primary side, and (iv) an arched valve surface. To explain these findings, two models were proposed. The first suggested limited lateral diatomaceous transport of Navicula species between the Galápagos and continental South America. Alternatively, these new species may be ecological specialists arising from the unique environmental conditions of the Galápagos coastal lagoons, which restrict the colonization of common diatom taxa and enable the establishment of novel, rare species. The Diablas wetlands are an important site for diatom research, where local‐scale environmental changes have combined with global‐scale biogeographic processes resulting in unique diatom assemblages.     

THE EVOLUTION OF ELONGATE SHAPE IN DIATOMS1

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.     

CYTOPLASMIC MASSES PRESERVED IN EARLY HOLOCENE DIATOMS: A POSSIBLE TAPHONOMIC PROCESS AND ITS PALEO‐ECOLOGICAL IMPLICATIONS1

In Lake Suigetsu, central Japan, greenish/light‐brown granules identified as cytoplasmic masses had been preserved in siliceous cell walls of freshwater diatoms in annual layers of lacustrine muds since the early Holocene. The lacustrine muds consisted of alternating dark‐colored (rich in diatom valves, clay, and organic matter) and light‐colored (mainly diatom valves) laminae. The greenish/light‐brown granules were predominately preserved in frustules of the genus Aulacoseira preserved in the dark‐colored laminae. The dark‐colored laminae were inferred to have formed annually under stratified water caused by surface water warming in summer that caused the formation of an organic‐rich anoxic layer on the lake bottom that favored granule preservation. The good preservation of cytoplasmic masses in dark‐colored laminae suggested a cause for diatom assemblage periodicity, a phenomenon that was commonly noted in temperate lakes: the cells containing these masses could be potential seed stocks for subsequent spring blooms. Frustules of the most abundant granule‐containing species, Aulacoseira nipponica (Skvortzow) Tuji, in the dark‐colored laminae of the Early Holocene muds were abundant in the overlying light‐colored laminae, suggesting that these species reproduced abundantly in springtime yielding a massive diatom bloom.     

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.     

SIZE DEPENDENCE OF GROWTH RATE,RESPIRATORY ELECTRON TRANSPORT SYSTEM ACTIVITY,AND CHEMICAL COMPOSITION IN MARINE DIATOMS IN THE LABORATORY1

The dependence of growth, electron transport system activity and chemical composition on the size of diatoms was examined during the exponential phase of growth. The six different marine centric species compared ranged in volume from 7.7 μm³ to 62 × 10⁵μm³. A size dependence was observed for growth, ¹⁴C uptake, respiration and the productivity index (¹⁴C/chl a). Although the size dependence of all parameters was similar, the results indicate that on a carbon basis, growth efficiency decreases with increasing size. The C/N and C/chl a ratios were not size dependent. The importance of the surface area to cell volume ratio, and the importance of carbon per unit volume in determining the observed size dependence are discussed.     

DAILY FLUCTUATIONS OF EXOPOLYMERS IN CULTURES OF THE BENTHIC DIATOMS CYLINDROTHECA CLOSTERIUM AND NITZSCHIA SP. (BACILLARIOPHYCEAE)1

Dynamics in the production of extracellular polymeric substances (EPS) were investigated for the benthic diatoms Cylindrotheca closterium (Ehrenberg) and Nitzschia sp. The effect of growth phase and light:dark conditions were examined using axenic cultures. Two EPS fractions were distinguished. Soluble EPS was recovered from the culture supernatant and represented polysaccharides that were only loosely associated with the cells. Bound EPS was extracted from the cells using warm (30° C) water and was more closely associated with the diatom aggregates. Concentrations of EPS exceeded internal concentrations of sugar throughout growth, indicating that EPS production is important in these organisms. Soluble and bound EPS revealed distinct differences in daily dynamics during the course of growth. Soluble EPS was produced continuously once cultures entered the stationary phase. During the stationary phase, chl a‐normalized EPS production rates equaled 6.4 and 3.4 d ^{? 1} for C. closterium and Nitzschia sp., respectively. In contrast, production of bound EPS occurred only in the light and was highest during the exponential phase. Up to 90% of the attached EPS that was produced in the light was degraded during the subsequent dark period. The monosaccharide distribution of EPS was constant during the course of the experiment. The soluble EPS consisted of high amounts of galactose and glucuronic acid, relative to rhamnose, glucose, xylose/mannose, and galacturonic acid. In contrast, glucose was the dominant monosaccharide present in the bound EPS. These differences suggest that the production of the two distinct EPS fractions is under different metabolic controls and probably serves different cellular functions.     

DISTRIBUTION OF EDAPHIC DIATOMS IN A MISSISSIPPI SALT MARSH: A CANONICAL CORRELATION ANALYSIS1

A canonical correlation analysis was performed to examine the degree to which edaphic diatom distribution in Graveline Bay Marsh, Mississippi could be related to environmental factors. It was possible to collapse the complex data base into two interpretable, orthogonal dimensions and to identify five distinct diatom communities. Diatom distribution on the marsh was primarily regulated by elevation and height of the overstory spermatophyte canopy. Although redundancy in the species data given the environmental data was relatively low (23% for the first two canonical variables), similar values were obtained for studies of benthic and planktonic diatoms in Yaquina Estuary, Oregon. The present study represents the first application of multivariate statistics to the study of salt marsh diatoms and demonstrates its usefulness in an environment where gradients are difficult to perceive.     

GROWTH CHARACTERISTICS AND ANTIOXIDANT PROPERTIES OF THE BENTHIC DIATOM NAVICULA INCERTA (BACILLARIOPHYCEAE) FROM JEJU ISLAND,KOREA1

Benthic diatoms are a commonly used food source in shellfish aquaculture. Diatoms of the genus Navicula are the most abundant benthic diatoms occurring year‐round on the coast of Jeju Island, Korea. We isolated an axenic strain of N. incerta Grunow; estimated its growth characteristics under 27 different combinations of temperature, salinity, and nutrients; and determined its biochemical composition and antioxidant activities. The maximum specific growth rate (μ_max), defined as the increase in cell density per unit time, was 0.81–1.04 · d^?1, and the maximum cell density, 7.99 × 10⁵ cells · mL^?1, was reached at 0.88 · d^?1 μ_max, 20°C, 30 psu salinity, and F/2 nutrient concentration on day 12 of the culture period. The approximate cellular composition was as follows: 7.0 ± 0.04% protein, 1.7 ± 0.28% lipid, 12.8 ± 0.85% carbohydrate, 68.4 ± 0.09% ash, and 10.1 ± 0.44% moisture. The antioxidant properties of N. incerta were determined for various extracts. The rates of 1,1‐diphenyl‐2‐picrylhydrazyl (DPPH) free‐radical scavenging for Neutrase and methanol extracts were 81.6% and 62.8%, respectively. Flavourzyme extract had a superoxide‐scavenging rate of 57.7%. Kojizyme and Ultraflo extracts had nitric‐oxide‐scavenging rates of 42.2% and 40.6%, respectively, significantly higher than commercial antioxidants, such as α‐tocopherol and butylated hydroxytoluene (BHT). The metal‐chelating activities of the methanol, Neutrase, and Termamyl extracts were 68.5%, 45.2%, and 41.2%, respectively, four to six times higher than commercial antioxidants. The Termamyl extract showed the highest linoleic acid peroxidation inhibition, exceeding α‐tocopherol and on par with BHT.     

PHOSPHATE AND SILICATE GROWTH AND UPTAKE KINETICS OF THE DIATOMS ASTERIONELLA FORMOSA AND CYCLOTELLA MENEGHINIANA IN BATCH AND SEMICONTINUOUS CULTURE1

Information on the nutrient kinetics of Asterionella formosa Hass. and Cyclotella meneghiniana Kutz. under either phosphate or silicate limitation was obtained for use in a Monod model and in a variable internal stores model of growth. Short-term batch culture growth experiments were fit to the Monod model and long-term semicontinuous culture experiments and short-term uptake experiments were fit to the variable internal stores model. Mathematical analysis indicates that the parameters of the 2 models may be expressed in terms of each other at steady state. The qualitative results of both batch and steady state culture methods agree. For limiting phosphate experiments. A. formosa is better able to grow at low PO₄-P concentrations than C. meneghiniana, as shown by its lower K for PO₄-P limited growth. The k_Q of A. formosa compared to C. meneghiniana found in long-term semicontinuous culture indicates that A. formosa is almost an order of magnitude more efficient at using internal phosphate for growth. The qualitative results under silicate-limited growth of C. meneghiniana is less than that of A. formosa. The k_Q from semicontinuous culture experiments indicates that C. meneghiniana is the more efficient at using internal silicate for growth. Nutrient uptake experiments showed more variability from a Michaelis-Menten relationship than short-term growth experiments. There were no significant differences between the 2 species in half saturation constants for either phosphate or silicate uptake. We observed a marked dependence of the coefficient of luxury consumption (R) of phosphate on the steady state growth rate. A. formosa has a higher R than C. meneghiniana.     

A PRELIMINARY INVESTIGATION OF THE PHYLOGENETIC RELATIONSHIPS AMONG THE FRESHWATER,APICAL PORE FIELD-BEARING CYMBELLOID AND GOMPHONEMOID DIATOMS (BACILLARIOPHYCEAE)1

Relationships among the apical pore field-bearing diatom genera Brebissonia, Cymbella, Gomphonema, Gomphoneis, Didymosphenia, Gomphocymbella, and Reimeria and their close ally Encyonema were evaluated using cladistics. These biraphid genera are a polythetic group defined by chloroplast number and type, mode of sexual reproduction, valve orientation and presence of apical pore fields. Character states of valve as well as non-siliceous features were described and then polarized using the outgroup method, with the naviculoid genera Anomoeoneis and Placoneis serving as outgroups. The cladistic analysis suggests two groups of terminal taxa, corresponding to to a predominantly cymbelloid lineage and a gomphonemoid lineage. Didymosphenia, previously thought to be closely allied to gomphonemoid diatoms, is shown to be more closely related to Cymbella. Gomphonema appears to be without distinguishing characteristics. Relationships of the other genera are described. The systematic position of small, doubly-punctate Gomphonema species is discussed.